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Bionomics, Systematics, and Phylogeny 

of Lytta, a Genus of Blister Beetles 

(Coleoptera, Meloidae) 



RICHARD B. SELANDER 



Illinois biological monographs: Number 28 



THE UNIVERSITY OF ILLINOIS PRESS 
URBANA, 1960 



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Bionomics, Systematics, and Phylogeny 
of Lytta, a Genus of Blister Beetles 

(COLEOPTERA, MELOIDAE) 




Lytta magister, male (drawing by F. Pacheco) 



Bionomics, Systematics, and Phylogeny 

of Lytta, a Genus of Blister Beetles 

(Coleoptera, Meloidae) 



RICHARD B. SELANDER 



ILLINOIS BIOLOGICAL MONOGRAPHS: Number 28 



THE UNIVERSITY OF ILLINOIS PRESS 
URBANA, 1960 



Board of Editors: william r. horsfall, r. d. de moss, francis j. krui- 

DENIER, WILSON N. STEWART, AND AUBREY B. TAYLOR 



THIS MONOGRAPH IS A CONTRIBUTION FROM THE DEPARTMENT OF ENTO- 
MOLOGY, UNIVERSITY OF ILLINOIS. DISTRIBUTED: DECEMBER 30, 1960. 



(g) 1960 BY THE BOARD OF TRUSTEES OF THE UNIVERSITY OF ILLINOIS. 
MANUFACTURED IN THE UNITED STATES OF AMERICA. LIBRARY OF CONGRESS 
CATALOG CARD NO. 60-15932. 






Contents 

Page 

Introduction 1 

Historical Resume 2 

Bionomics 2 

Systematies 3 

Methods and Terms 4 

Acknowledgments 6 

Bionomics 8 

Economic Importance 9 

Damage to cultivated plants 9 

Medical importance 9 

Enemies 10 

Activities of Adults 11 

Seasonal distribution and longevity 11 

General behavior 12 

Courtship and mating 14 

Oviposition 15 

Development of Immature Stages 16 

Incubation 16 

Number of larval instars 16 

Larval hosts 17 

First larval instar 19 

Grub stage 19 

Coarctate larval instar 19 

Ultimate larval instar 20 

Pupal stage 20 

Systematics and Phylogeny 21 

Genus LYTTA Fabricius 21 

Limits of the Genus 22 

Classification 26 

Conspectus 26 

Basis 27 

Old World subgenera 27 

Key to Subgenera 28 

Phylogeny of the Subgenera 30 

Distribution 31 



Vi BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Page 

Artificial Key to the North American Species of Lytta 35 

Subgenus PARALYTTA, new subgenus 49 

Nitidicollis group 53 

Fulvipennis group 57 

Variabilis group 61 

Variabilis subgroup 62 

Koltzei subgroup 73 

Quadrimaculata subgroup 84 

Reticulata group 101 

Reticulata subgroup 102 

Deserticola subgroup 108 

Magister group 113 

Tenebrosa subgroup 115 

Melaena subgroup 122 

Magister subgroup 127 

Subgenus ADICOLYTTA, new subgenus 137 

Peninsularis group 141 

Eucera group 143 

Mutilata group 147 

Subgenus POREOSPASTA Horn, new status 157 

Stygica group 161 

Moerens group 193 

Moerens subgroup 194 

Cyanipennis subgroup 204 

Childi subgroup 215 

Subgenus POMPHOPOEA LeConte, new status 222 

Polita group 226 

Aenea group 228 

Sayi group 234 

Literature Cited in the Text 240 

Index 285 



Introduction 

The genus Lytta, with 128 currently assigned species, is one of the 
larger genera of the family Meloidae, which is a member of the section 
Heteromera of the superfamily Cucujoidea of the order Coleoptera. Adult 
Meloidae, known popularly as blister beetles, are rather slender, soft- 
bodied, long-legged, phytophagous insects. In the drier subtropical and 
tropical regions of the earth they form a very conspicuous part of the 
fauna, feeding gregariously, often in spectacular numbers, and at times 
causing severe damage to crops. In more mesic regions the family is not 
so well represented. In the larval stage Meloidae are unusual for Cole- 
optera in having a parasitic mode of life. Larval hosts are of limited 
variety: wild bees and grasshoppers. Meloid larvae begin life as active, 
strongly sclerotized campodeiform insects charged with the responsi- 
bility of locating their own food materials. In Lytta and a large number 
of related genera this is achieved by active searching; in other genera 
phoresy is involved. Developing larvae undergo hypermetamorphosis, 
passing through four morphologically and behaviorally different phases 
before reaching the pupal stage. 

My attention was first drawn to the genus Lytta in the summer of 1950 
when, while collecting insects at high elevation in the Wasatch Range 
of northern Utah, I found adults of the large, metallic greenish blue 
cyanipennis swarming on one of the native lupines. During this same sum- 
mer I obtained eggs from some of the adult beetles and subsequently 
succeeded in rearing larvae of the species through several instars. Later, 
working in Mexico, I had the opportunity to observe additional species 
of Lytta in their natural habitats and to secure additional larvae. In 
selecting the genus for study, I was influenced both by the experience 
I had had with the group in the field as well as by the knowledge that 
previously no attempt had been made to define the genus on a world basis, 
to incorporate both Old and New World species into a scheme of classi- 
fication or phylogeny, or to revise the species of an entire continent. 
Further, since the genus stands in the midst of the complex and un- 
wieldy tribe Lyttini, it seemed to me that a clarification of the limits 
and composition of the genus would be an especially important contri- 
bution to the study of the family. 

Within limits imposed by time and the availability of material, the 
present study is as comprehensive as it has been possible to make it. A 
major part is devoted to a revision of the species of Lytta of the North 

1 



2 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

American continent. In the phylogenetic portions of the study the fauna 
of North America is treated at the specific level, while consideration of 
the fauna of the Old World is, of necessity, confined to main evolution- 
ary lines. Similarly, the formal classification includes all the species of 
the genus in North America but only a part of the species in the Old 
World, i.e., those represented in collections studied or described rather 
fully in the literature. Species excluded from consideration for one reason 
or another are mentioned in the discussion of limits of the genus. 

Historical Resume 

Bionomics. A detailed review of the history of discovery of the larva 
and mode of larval life of Lytta was given by Beauregard (1890) in his 
treatise on the family Meloidae. 

According to Beauregard, the larva of Lytta was first discovered by 
Loschge, who in 1788 gave a short description of the first instar of 
vesicatoria. For the next three quarters of a century little of real conse- 
quence was published regarding the bionomics of the genus aside from 
scattered observations on mating behavior and other adult activities of 
vesicatoria, most of which are cited in the following pages under appro- 
priate subdivisions of the discussion. 

The association of the larval stages of Lytta with Hymenoptera was not 
established until after it had been shown that several other genera of 
Meloidae are larval parasites in the nests of bees. It was recognized, 
however, that larvae develop below the surface of the ground, for it 
became widely held among entomologists that the larvae of Lytta fed 
on the roots of plants. In 1875, Lichtenstein began a series of attempts to 
rear larvae of vesicatoria on the provisioned pollen of several species 
of bees. Reports of his progress appeared in numerous short notes that 
culminated in 1879 in the announcement of a completed rearing, using 
pollen of a species of Ceratina bee. Following this, Beauregard (1890) 
carried out additional rearings of vesicatoria, made further observations 
on the habits of larvae and adults, and described all immature stages in 
detail. His work is the last to contribute anything of significance to our 
knowledge of the bionomics of the genus Lytta in the Old World. 

In more recent times, Linsley and MacSwain ( 1942, 1952, and 1958 ) 
and Linsley, MacSwain, and Smith (1956) have made observations on 
some aspects of the life histories of the North American species Lytta 
melaena, tenebrosa (as occipitalis), chloris, stygica (as purpurescens), 
moerens, and variabilis. 

Data on bionomics contained in the published literature of Lytta are 
supplemented in the present work with information that I have recorded 
for the genus in the course of my study of the Meloidae over the past 
ten years. 



INTRODUCTION 



Systematics. In the New World the main systematic works dealing with 
species now assigned to the genus are Horn's ( 1873 ) reviews of Lytta 
and Pomphopoea for the United States, Fall's ( 1901 ) revision of Horn's 
keys to the species of Lytta, Champion's (1892) synopsis of Lytta for 
Mexico and Central America, and MacSwain's (1956) treatment of the 
first instar larvae of many of the species of Lytta and Pomphopoea of the 
United States. In the Old World the principal systematic works on 
the genus Lytta are Escherich's ( 1895 ) revision of some of the Palearctic 
species and Peringuey's (1909) revision of the South African species. 

Horn, and later Fall, divided the species of the genus Lytta of the 
United States into three groups, as follows: 

Group I. Antennae of male with intermediate joints (4-5-6-7) either de- 
formed or much thicker than those which precede or follow. Antennae more 
or less moniliform in both sexes, gradually stouter to tip in the female and 
rarely longer than the head and thorax, never equal to half the length of body. 
Type, C. vulnerata, Lee. 

Group II. Antennae not dissimilar in structure in the sexes, either moniliform 
or slender, always gradually stouter toward the tip, in several species longer 
than half the length of body. Types, C. cardinalis, Chev. (moniliform antennae) 
[misidentification of falvipennis LeConte], C. stygica, Lee. (slender antennae). 

Group III. Antennae with compressed joints; joints 5-10 usually broader than 
long, thorax very convex, somewhat spherical in form. Both hind tibial spurs 
slender and acute. Type, C. compressicornis, Horn. 

In its final form (Fall, 1901), Group I of this classification included, 
in addition to vulnerata (= cooperi), the species eucera, mutilata, pen- 
insularis, quadrimaculata, margarita, magister, deserticola, morosa, mel- 
aena, nuttalli, cyanipennis, viridana, tenebrosa, lugubris (= ulkei), 
childi, funerea, puberula, and biguttata. Group II included, in addition 
to fulvipennis and stygica (= stolida, purpurescens, and difficilis), the 
species suavissima (= gentilis), moerens ( — incommoda), insperata, 
molesta, reticulata, agrestis, cribrata, lecontei (= dichroa), occipitalis 
(= tenebrosa), chloris, nigripilis, auriculata, refulgens, aeneipennis, 
crotchi, nitidicollis, lugens, and rathvoni. The species of Group III, along 
with suavissima Wellman, were later transferred from Lytta to form the 
epicautine genus Linsleya MacSwain ( 1951 ) . 

As constituted, both Groups I and II of Horn and Fall's classification 
are polyphyletic. In the main this is due to the fact that the incrassate 
condition of the intermediate male antennal segments characteristic of 
many of the North American species of Lytta has developed independ- 
ently in two entirely distinct phylogenetic lines. In addition, both authors 
erred in their interpretation of this character. Actually, male and female 
antennae are to some extent dissimilar in all species of the genus Lytta. 
Further, the intermediate male antennal segments in some species, e.g., 
fulvipennis, although definitely incrassate, are no wider or only slightly 



4 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

wider than the segments that precede and follow them. Again, the degree 
of enlargement of the intermediate segments may vary considerably 
within a single species. The highly subjective nature of the group criteria 
employed by Horn and Fall is shown dramatically by the fact that they 
placed tenebrosa and occipitalis in different groups, although these forms 
are in reality only geographical variants of the same species. 

Champion did not propose a classification of the species he treated. 
In the selection of characters he tended to follow Horn. His key to species 
is incomplete, and the distinctions he made between species are in some 
cases inexact. 

MacSwain proposed a division of the species of Lytta of the United 
States into four groups, one of which ( Nitidicollis Group) he regarded 
as probably unnatural. Fundamentally, my studies of adult beetles have 
confirmed his conclusions concerning the systematics of the genus based 
on larval morphology ( but not necessarily those based on adult morphol- 
ogy). As a result, I have adopted his concept of a Magister Group, 
Moerens Group, and Polita Group (herein called Stygica Group). 

Methods and Terms 

The classification proposed in the present work is a phylogenetic one. 
In essence this means that, so far as possible, the criterion for the forma- 
tion of taxa is not degree of similarity or difference of the species but 
their phylogenetic relationships. Phylogeny as the theoretical basis for 
systematics has recently received a good deal of criticism from one group 
of systematists, but Simpson (1959) has, I think, answered their argu- 
ments. In preparing phylogenies for this study, much use has been 
made of the principle that the correct arrangement of species and higher 
categories is most likely to be the one that accounts for the origin and 
distribution of characters in the simplest manner. In many cases it has 
been possible to deduce with some confidence the primitive characters 
of taxa and then to trace the evolutionary lines from a hypothetical 
ancestral type. In others, it has been necessary to regard degree of 
morphological similarity (static relationship of Michener and Sokol, 
1957) as indicative of phylogenetic relationship. In actual practice this 
last approach does not differ from the nonphylogenetic approach to 
systematics, although from a theoretical viewpoint the distinction between 
the two is a critical one. 

About one-fourth of the North American species of Lytta show marked 
geographic variation. In dealing with species in which only a single 
character varies geographically or in which variation of characters is 
geographically discordant, I have simply described the variation observed. 
Where species exhibit concordant geographic variation, the concept of 
subspecies has proved convenient. In line with views expressed by Wilson 



INTRODUCTION 5 

and Brown (1953), however, vernacular names have been used for the 
subspecies recognized here. 

In mapping the geographic distribution of species of Lytta, definite 
locality records have been indicated by solid figures and indefinite ones, 
such as state records, by open figures. 

Some comment about the keys in this work is in order. As far as prac- 
ticable, the keys to subgenera, groups, subgroups, and species have been 
made phylogenetic, with the intention of their serving as convenient 
summaries of the morphological basis for the classification. Their ease of 
use in the identification of specimens has therefore been a minor con- 
sideration. Males will probably run in them satisfactorily, but in most 
cases thev will not serve for the identification of females unless the 
group or subgroup to which a species belongs is already known. For the 
identification of females and (in many cases) more convenient identifica- 
tion of males, the artificial key beginning on page 35 should be used. 

There are only a few morphological terms that require explanation. 
Vertex, as used in this work, refers to that part of the head capsule above 
the dorsal margin of the eyes. In describing the male genitalia, Michener's 
( 1944 ) suggestion that the terms gonoforceps, gonostylus, and aedeagus 
be used in place of the terms tegmen, lateral lobe, and median lobe, 
respectively, has been followed. The term gonocoxal plate replaces the 
term basal piece. Reference to abdominal sterna is always to externally 
visible sterna. The fifth and sixth sterna are morphologically the seventh 
and eighth, respectively. 

Antennal length is usually expressed in this work in terms of the point 
that the antennae reach relative to the pronotum. In determining this 
point the antennae are directed over the vertex and then posteriad along 
the dorsum of the body of the specimen. In a few cases, where more 
critical measurements are desirable, absolute antennal length is com- 
pared with fore tibial length. 

Synonymies given for the species are complete except for references 
given in catalogs. For the spelling of place names the following works 
were consulted: Rand McNally Road Atlas for Canada and the United 
States; Gazetter No. 15, Mexico, of the United States Roard on Geo- 
graphic Names for Mexico; and the current National Geographic Society 
Map of Mexico and Central America for Central America. Certain place 
names in Mexico, such as San Luis Potosi, may refer to both a state and 
the capital city of the state. These are always interpreted in this work 
as state names, unless there is evidence to the contrary. 

In the citation of bibliographic references, the system of abbreviation 
of names of journals contained in the Style Sheet for Scientific Publica- 
tions of The American Museum of Natural History has been followed. 
In the text authors' names are not cited for taxa that are revised in this 



6 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

study inasmuch as this and other bibliographic information is given 
in connection with the individual treatment of these taxa. For all other 
taxa authors' names are cited in the text at least once. 

Acknowledgments 

In the course of this study field work in Mexico and the southwestern 
United States was supported in part by research fellowships from the 
National Science Foundation (1953) and the University of Illinois (1952) 
and grants from the University of Utah ( 1954 ) , the Penrose Fund of the 
American Philosophical Society (1954), and the Sigma Xi-RESA Re- 
search Fund ( 1954 and 1955 ) . I am happy to express my appreciation to 
these institutions for their assistance. In addition, I would like to thank 
the University of Illinois Research Board for a grant covering the cost 
of typing the final draft of the manuscript. 

For the loan of specimens of Lytta from institutional collections now 
or formerly in their charge, I am indebted to the following colleagues: 

C. P. Alexander, University of Massachusetts; W. V. Balduf, University 
of Illinois; Douglas Barnes and William W. Gibson, Oficina de Estudios 
Especiales, Programa Cooperativo de Agricultura entre la Secretaria y 
la Fundacion Rockefeller, Mexico, D. F., Mexico; William F. Barr, 
University of Idaho; Osmond P. Breland, University of Texas; George 

D. Butler, Jr., and Floyd G. Werner, University of Arizona; George W. 
Byers, University of Kansas; W. J. Brown and Henry F. Howden, 
Canadian National Collection; Leland Chandler, Purdue University; 
P. J. Darlington, Jr., Museum of Comparative Zoology, Harvard Univer- 
sity; Henry Dietrich, Cornell University; Lawrence S. Dillon, Agricul- 
tural and Mechanical College of Texas; Henry S. Dybas and Rupert L. 
Wenzel, Chicago Natural History Museum; George F. Edmunds, Jr., 
University of Utah; Wilbur R. Enns, University of Missouri; Heinz 
Freude, Zoologische Staatssammlung, Munich; Harold J. Grant, Jr., and 
James A. G. Rehn, Philadelphia Academy of Natural Sciences; Christine 
M. F. von Hayek, British Museum (Natural History); C. Clayton Hoff, 
University of New Mexico; Paul D. Hurd, Jr., and J. W. MacSwain, Uni- 
versity of California, Berkeley; Andre Janssens, Institut Royal des 
Sciences Naturelles de Belgique; George F. Knowlton, Utah State Uni- 
versity; Ira LaRivers, University of Nevada; Hugh B. Leech, California 
Academy of Sciences; A. T. McClay, University of California, Davis; 
Raul MacGregor, Direccion General de Defensa Agricola, Mexico, 
D. F., Mexico; Jean M. Mathieu, Instituto Tecnologico y de Estudios 
Superiores de Monterrey; T. E. Moore, University of Michigan; Vincent 
D. Roth, Oregon State College; M. W. Sanderson, Illinois Natural History 
Survey; Joseph C. Schaffner, Iowa State College; T. J. Spilman and 
George B. Vogt, U.S. National Museum; J. W. Tilden, San Jose State Col- 



INTRODUCTION 7 

lege; Fred Truxal, Los Angeles County Museum; Patricia Vaurie, Ameri- 
can Museum of Natural History; Leonila Vazquez, Instituto de Biologia, 
Universidad de Mexico; George Wallace, Carnegie Museum; John A. 
Wilcox, New York State Museum; Stephen L. Wood and Vasco M. Tan- 
ner, Brigham Young University. Messrs. Freude, MacSwain, Leech, and 
McClay were especially helpful in providing me with material and in- 
formation, as was Miss von Hayek, who also made drawings of type 
material in the British Museum (Natural History) for my use. 

Specimens of Lytta from their private collections were generously 
made available for study by George E. Bohart, Candido Bolivar, R. H. 
Crandall, Henry F. Howden, Frank H. Parker, John H. Robinson, and 
Floyd G. Werner. Gifts of specimens collected by them were received 
from F. F. Bibby, John K. Bouseman, Harold R. Dodge, John C. Downey, 
David R. Lauck, Robert K. and Bonnie J. Selander, Philip W. Smith, 
and Barry D. Valentine. The co-operation of all these individuals is 
greatly appreciated. 

For their interest and assistance in my work over the past several years 
I am indebted to H. H. Ross, under whose direction this study was ini- 
tiated, and to M. W. Sanderson. To Bill M. Woods, formerly in 
charge of the Map and Geography Library of the University of Illinois, 
I owe a great debt of gratitude for the many hours he spent in tracing 
down obscure localities for me. Finally, I want to acknowledge the in- 
valuable assistance I received from Jolene M. Flamand throughout the 
course of the present study, both in the laboratory and in the field. 



Bionomics 

This section is a summary of available information concerning the 
development, habits, and ecological relationships of the species of the 
genus Lytta. As will become evident, this information is fragmentary. 
For a large number of species of the genus even such basic data as food 
plant records are lacking, and our knowledge of details of larval develop- 
ment is based in great part on observations of a single species (vesi- 
catoria ) . 

So far as it is known, there are only two orders of insects that provide 
larval food material for Meloidae. Members of the meloine tribes Epi- 
cautini and Mylabrini feed on the eggs of grasshoppers; all other Me- 
loidae parasitize the nests of wild bees, feeding on provisions and (in 
many cases) immature stages of the bees. On the basis of the phylo- 
genetic position of the tribes Epicautini and Mylabrini indicated by 
studies of morphology, it seems reasonable to conclude that feeding on 
the eggs of grasshoppers is a specialized characteristic within the Me- 
loidae. Interestingly, no major adaptive modification seems to have been 
necessary to permit the evolutionary change from parasitism of bees 
to parasitism of grasshoppers, for the general features of bionomics are 
similar for all Meloidae. 

With the exception of a few degenerate species, e.g., members of the 
genera Hornia Riley and Tricrania LeConte, that do not leave the vicinity 
of their site of larval development, all Meloidae are phytophagous in 
the adult stage. Some species eat only pollen or leaves; others feed more 
generally. Members of the subfamily Meloinae oviposit in burrows in 
the ground excavated by the female beetles. Members of the subfamily 
Nemognathinae usually oviposit on their food plants. The only exception 
to this last rule occurs in the degenerate forms mentioned above. Larval 
development in all cases is hypermetamorphic, i.e., there are striking 
morphological differences between larvae of several of the instars. Larvae 
in the first instar invariably are active, strongly sclerotized forms. They 
actively seek out their food in all Meloinae except Meloe Linnaeus. In 
Meloe and in all Nemognathinae they attach to adult bees and are 
passively carried to their food. Once the larvae have begun feeding, they 
pass through three or four grublike instars. After feeding has been 
completed, they enter the inactive coarctate instar in which the append- 
ages become rudimentary. A return to a grublike form takes place in 
the next instar, which is followed shortly by pupation and emergence of 
the adult beetle. 



BIONOMICS 



Economic Importance 



Damage to cultivated plants. Although a few species of Lijtta feed on 
cultivated plants, their attacks are of such a sporadic and local nature 
that they are not of major economic importance. In the eastern United 
States Lytta (Pomphopoea) aenea, polita, and saiji have been reported 
as pests of peaches, plums, apples, and pears. The adult beetles feed on 
the flowers and young fruit, at times causing severe local damage to 
commercial plantings. Three North American species of Lytta have been 
recorded from field crops. These are moerens, reported once from 
radishes; nuttalli, from beans, beets, oats, and barley; and cyanipennis, 
from beans and peas. Some of these records are probably based on acci- 
dental associations. In Europe feeding aggregations of adults of vesi- 
catoria sometimes defoliate ornamental shrubs. 

Medical importance. From an economic standpoint Lytta has long been 
of interest and importance as a source of cantharidin. This substance is 
an odorless, colorless anhydride found in all Meloidae with the possible 
exception of the species of the tribe Horiini. Its physical and chemical 
properties are discussed by Fumouze (1867) and Gornitz (1937). It has 
been obtained commercially from a number of species of Meloidae 
(particularly members of the genera Mylabris Fabricius and Epicauta 
Dejean), but by far the most important source has been Lytta vesica- 
toria, commonly known as the Spanish fly (mouche d'Espagne). 

According to Beauregard ( 1890 ) , cantharidin is produced in the third 
pair of seminal vesicles of the male and in the copulatory vesicle and 
ovaries of the female. It is distributed to the blood and other soft 
tissues of the body but is not incorporated into the exoskeleton. It has 
been shown that it is also present in the eggs and first instar larvae. 
Its function in the Meloidae is unknown. The characteristic odor of 
Meloidae is said to be produced not by cantharidin or any of its deriva- 
tives but by an unidentified essential oil ( Fumouze, 1867 ) . 

For commercial purposes cantharidin is seldom used in its pure form; 
rather, the dried beetles are ground up to form a crude drug known as 
cantharides. The method of collection of adults of Lytta vesicatoria and 
their preparation as cantharides were described by Fumouze (1867). 
In the morning, before the beetles have recovered from the effects of 
the coldness of the night, large sheets of cloth are spread at the bases of 
trees upon which they feed. The trees are then shaken, and the beetles 
tumble onto the cloths. Beetles collected in this manner are killed in 
boiling water or hot vinegar and dried in the air or in ovens. 

During the past century most of the cantharides used in Europe and 
the United States has come from eastern Europe and Italy. Considerable 



10 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lyttd 

variation of cantharidin content in samples of vesicatoria from different 
parts of Europe was reported by Fumouze (1867), with a range from 
1.7 per cent cantharidin in a German sample (which Fumouze felt was 
probably adulterated) to 5.0 per cent in a sample from Trieste. 

When administered to humans, cantharidin is an active irritant and 
vesicant. On the skin it produces "redness, smarting, and pain, followed 
very soon by small vesicles, which later coalesce into one large blister" 
(Cushny, 1901, p. 89). In the formation of the blister the outer layers 
of the epithelium separate from the corium, the cavity produced being 
filled with a yellow serous fluid containing leucocytes (Meyer and Gott- 
lieb, 1926). Taken internally in large quantities cantharidin produces 
vesication of the mucous membranes, vomiting, purging, abdominal pain, 
and shock. Comparatively small quantities irritate the bladder, causing 
a constant desire to urinate. In women it reportedly can produce abortion, 
while in both sexes irritation of the urinary tract may lead to increased 
sexual desire. 

In contrast to humans, some animals (chicken, hedgehog, and frog) 
reportedly have a high degree of immunity to cantharidin (see Cushny, 
1901). 

The history of the use of cantharidin can be traced in detail in the 
works of Fumouze (1867), Beauregard (1890), Escherich (1895), and 
Gornitz (1937). The ancient Greeks and Romans employed it as an 
aphrodisiac, poison, and medicinal drug. Its value as an aphrodisiac for 
humans has been disputed, but it seems to have been used as such until 
relatively recently and is still used in some countries in the breeding of 
cattle. It occupied a prominent position in heroic medicine, and until 
the present century it was commonly administered orally as a remedy 
for a variety of maladies, including tuberculosis and urinary disorders. 
At present its internal use has been largely discontinued, although it 
continues to enjoy some popularity as an irritant and counterirritant ap- 
plied to the skin. For this purpose it is commonly used in the form of a 
"cantharides plaster" containing 0.1 gram of cantharides per square 
centimeter (Goodman and Gilman, 1941). According to Cushny (1901, 
p. 91), it has also been used as "a constituent of hair washes, its irritant 
action on the skin being credited with causing a more rapid growth of 
hair." 

When applied to certain insects cantharidin acts as an effective nerve 
poison. The possibility of its use as an insecticide in this connection 
has been investigated by Gornitz (1937). 

Enemies 

Adult meloids are known to be attacked by nematodes, spiders, Hemip- 
tera, ants, and ceratapogonid flies. There are, however, few specific 



BIONOMICS 11 

records of invertebrate predation, and only one of these involves a species 
of Lytta. This is Lytta nuttalli, which is preyed on by the mirid bug 
Hadronema militaris Uhler. Similarly, there is little specific information 
available regarding predation on meloids by vertebrates. A number of 
field records of birds, lizards, and toads eating members of the genus 
Epicauta have been reported, but there are no records of meloid predation 
by mammals. Marshall (1902), in Africa, found that several species 
of Mylabrini and single species each of Eletica Lacordaire and Zonitis 
Fabricius were consistently refused as food items by captive baboons 
and a falcon. Carpenter ( 1921 ) reported that two Cercopithecus monkeys 
whose food habits he studied in Africa did not eat meloids, although one 
of them tasted specimens. Pritchett (1903) reported negative results in 
her attempts to feed adults of Lytta fulvipennis to caged Sceloporus and 
Gerrhonotus lizards in Texas. According to Pritchett's account, the lizards 
seized three of several of the meloids offered but quickly rejected them. 
After briefly chewing a meloid, one of the Gerrhonotus lizards "began 
writhing and rubbing his mouth in the sand, appearing much distressed" 
(p. 279). 

As larvae, species of Lytta and other Meloidae are subject to attack by 
a number of parasites and predators associated with their larval food 
materials. So far as it is known, however, none of these specifically 
attacks Meloidae. In this connection, it has been suggested by Linsley 
and MacSwain ( 1952 ) that in Lytta the larva's habit of burrowing away 
from the cell series in which it develops may serve as a means of protec- 
tion against attack by enemies. 

Activities of Adults 

Seasonal distribution and longevity. Data on the seasonal distribution 
of adults are available for 63 of the 68 species of Lytta occurring in 
North America. Known periods of activity of adults range from one 
month (for 11 species) to ten months (for magister and vulnerata). The 
average length of the period in North America is three and one-half 
months. The period of activity of adults of vesicatoria in France is re- 
ported to last two and one-half months (Beauregard, 1890). 

For the majority of species of Lytta the period of activity of adults 
falls in middle and late summer. However, in the subgenus Poreospasta 
it falls primarily in spring and early summer. Of the 21 species of this 
subgenus, only 3 (nuttalli, cyanipennis, and viridana) are recorded in 
the adult stage as late as August and onlv 1 (nuttalli) as late as Septem- 
ber. Only three North American species of Lytta appear to be active as 
adults through the winter months. These are polita (recorded from mid- 
December to early June), scituloides (apparently active from mid- 
September to mid-May), and eucera (recorded in February and in ever} 7 



12 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LljttCl 

month from May to December). There is no definite indication of more 
than one generation per year for any species of the genus. 

It is highly doubtful that individual beetles live for the entire period 
of activity of adults of most species. Adults of vesicatoria live about four 
weeks under laboratory conditions (Beauregard, 1890). I have kept 
adults of several species of Lytta alive in captivity for periods of two to 
three weeks but have never determined how long it would be possible to 
maintain them. In all probability, adults of most species do not live much 
longer than those of vesicatoria. Consequently, emergence of adults of 
most species must take place over a period of several weeks. Beauregard 
noted that in France a first emergence of vesicatoria occurs in mid-May, 
followed by a second in mid- June, and a possible third a little later. This 
same author felt that this prolongation of time of emergence reflects 
differences in exposure of terrain in which the larvae develop as well as 
the fact that larvae remaining dormant over more than one winter attain 
the adult stage and emerge from the ground earlier than do individuals 
overwintering only a single year. Differences in latitude and elevation as 
well as annual climatic variation undoubtedly exert a great influence on 
the time of emergence of adult beetles of many species of Lytta. Never- 
theless, it seems reasonable to suppose that the seasonal distribution 
of a given species is synchronized very closely with the period of nesting 
activity of its larval host or hosts. 

General behavior. The gregarious habit of adults of the genus Lytta 
and many other genera of Meloidae probably functions in part as a 
mechanism to keep the beetles near nesting sites of host bees and thus 
to assure the larvae of a reasonable chance of finding their way into 
suitable bee cells. 

Compared with adults of Epicauta and Pyrota, those of Lytta are for 
the most part slow-moving and unwary. Frequently they can be ap- 
proached to within a distance of a few inches before they become dis- 
turbed. When disturbed they usually remain motionless or attempt to 
crawl away from the disturbing object; only rarely do they drop from 
their positions or take flight. They feign death and undergo reflex 
bleeding only when handled roughly. Some differences in general be- 
havior exist between species. For example, adults of ebenina and sangui- 
nea are more active and alert than those of such species as quadrimacu- 
lata, eucera, and cyanipennis. 

Adults of most species are more active during the day than thev are 
during the night. In some cases, adults spend the night on their food 
plants; in others, they crawl to the base of the plants and remain at 
ground level until morning. They are not attracted to light at night. 

In the course of the present study the feeding habits of adults were 
observed at first hand for the species variabilis, michoacanae , ebenina, 



BIONOMICS 13 

plumbea, biguttata, scitula, quadrimaculata, eucera, sanguined, mutilata, 
and cyanipennis. Adults of mutilata eat the entire body of their food 
plants, which are members of the parasitic plant genus Cuscuta. Adults 
of eucera and cyanipennis feed primarily on leaves, although in both 
species flowers are also eaten and seem to be preferred. The feeding 
habits of adults of vesicatoria are presumably similar to those of these 
last two species. Adults of the other species listed above apparently feed 
only on flowers ( pollen and petals ) . From personal observation and from 
data found in the literature and on the labels of specimens examined, 
the following tentative generalizations may be made regarding the feed- 
ing habits of the genus. In the subgenera Paralytta and Pomphopoea and 
in all but one or two groups of Poreospasta, adults feed only on flowers. 
In the Cyanipennis Group (and possibly the Moerens Group) of Poreo- 
spasta and in the subgenus Lytta they feed on both flowers and leaves. 
In Adicolytta they may be restricted to flowers, e.g., sanguinea, or may 
eat both flowers and leaves, e.g., eucera and mutilata. 

Food plant records for the North American species of Lytta are given 
in the following section of this work. Where the species of a subgenus 
or group exhibit marked preference for plants of one or a few families, 
this fact is mentioned in the discussion of the appropriate taxon. The 
families of plants which are of most importance as sources of food for 
adults of Lytta in North America are Leguminosae, Convolvulaceae, 
Compositae, Papaveraceae, and Rosaceae. In the Old World food plant 
records are available for only two species, both belonging to the subgenus 
Lytta. These are caraganae, which feeds on Caragana (Leguminosae), 
and vesicatoria, which has been recorded from plants of the families 
Oleaceae (Fraxinus, Liqustrum, Olea, and Syringa), Caprifoliaceae 
(Lonicera, Sambucus, and Symphoricarpos), and Salicaceae (Populus 
and Salix) (with some unverified records of attacks on cereals and other 
grasses) ( Lichtenstein, 1877; Beauregard, 1890; Lampa, 1903; and Houl- 
bert and Betis, 1913). 

In general, few if any species of Lytta are confined to a single species 
of plant. Some species seem to have a strong preference for representa- 
tives of a single genus of plants, while perhaps the majority show such a 
preference for members of a single family. Species that ordinarily feed 
on only one genus or family of plants can utilize other plants, at least 
as a dietary supplement. For example, quadrimaculata feeds only on 
Papaveraceae when it is available but in its absence will turn to Com- 
positae. 

There is evidence that some species of Lytta feed on the same kinds 
of plants that their bee hosts utilize as pollen sources. The advantage to 
the meloid of this relationship seems obvious, although the extent to 
which it is realized is uncertain. 



14 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Courtship and mating. Courtship in Lytta consists basically of the male 
talcing a position directly above the female and attempting to stimulate 
her to accept genital contact. Stimulation seems to be derived principally 
from stroking, flagellation, or other manipulation of the female's antennae 
by the male's antennae and (in some cases) fore legs. When genital con- 
tact is achieved, the male dismounts and turns to face in the opposite 
direction from the female, in the usual mating position for species of 
the subfamily Meloinae. 

Males frequently have the antennae and legs specially adapted for use 
in courtship activity. In Paralytta, Adicolytta, and a few species of Poreo- 
spasta the intermediate segments of the male antennae are enlarged, 
while in some species of the second subgenus they are further modified to 
form a clasping mechanism. Adaptations of the male legs are varied. In 
most species of the genus the tarsal pads of the male are larger and 
denser than those of the female. In Adicolytta the femora and tibiae of 
the fore and middle legs may be distorted. In Poreospasta the hind tro- 
chanters are sometimes spined or angulate, and the tibial spurs and first 
tarsal segment of the fore legs are frequently modified. Adaptations of 
this last sort are also found on the fore legs of species of the subgenus 
Lytta and on both the fore and middle legs of the Pseudolytta. In males 
of all species of the genus the last visible abdominal sternum is emargi- 
nate, which permits the aedeagus to be thrust forward during courtship. 

Courtship behavior in Lytta vesicatoria was described by Goedart 
(1700), Audouin (1826), Fabre (1886), and Beauregard (1890). In this 
species the male begins courtship by gently stroking the under surface 
of the female's thorax with his fore and middle legs. Then the antennae 
are used to flagellate the head of the female, and the abdomen (which 
is elongated) is vibrated very rapidly as the male attempts to make 
genital contact. Periodically the male grasps the antennae of the female 
with his fore legs and pulls vigorously on them. A socket formed by the 
first tarsal segment and tibial spur on each fore leg of the male provides 
a device for holding the antennae of the female. 

I have observed courtship activity in Lytta cyanipennis and variabilis. 
In cyanipennis the male extends the antennae forward and strokes the 
antennae of the female; at the same time he violently shakes his entire 
body and vibrates his abdomen (which is elongated) so rapidly that 
it is blurred. At one time a male under observation continued this be- 
havior, with short periods of rest, for five hours. In variabilis the male 
uses his antennae to draw back the antennae of the female so that the 
second to fourth segments of first one and then the other of them are 
pressed into the finely punctulate, short-pubescent concavity on each 
side of his head. This alternate manipulation of the female antennae 
is repeated at intervals of about one second. The female sometimes 



BIONOMICS 15 

raises her antennae in response, facilitating the action of the male. The 
female may be stimulated by a chemical substance in the modified 
areas of the head of the male or by contact with the short setae present. 
Modifications of the head of the type found in the male of variabilis 
are characteristic of some 22 species forming the Variabilis and Reticulata 
groups of the subgenus Paralytta. 

Mating in the genus Lytta, as in other Meloinae, generally lasts a long 
time. Pairs of adults of vesicatoria have remained coupled from 4 to 20 
hours ( Beauregard, 1890 ) ; a mating of cyanipennis observed by me lasted 
IP/2 hours. Beauregard's (1890) suggestion that actual physiological 
mating or insemination is completed in a relatively short time is probably 
correct, although this has not been established definitely. 

Adults continue to feed while coupled. In the course of their activities 
they frequently pull in opposite directions or hang from each other. 
However, their coupling mechanism is so effective that they seldom if 
ever lose contact accidentally. In fact, mating pairs sometimes experience 
difficulty in disengaging. In some cases the aedeagus of the male may 
even be pulled from his body. 

Two structural adaptations of the genitalia are basic to the coupling 
mechanism. One is the presence of hooks on the aedeagus of the male. 
These provide for secure attachment by catching on folds of the vagina 
of the female. The other adaptation is the great reduction in length of 
the genital tube of the female. In most Coleoptera the genital tube is a 
long, membranous structure. In repose it is entirely concealed within 
the abdomen of the female; during oviposition it is extruded to its full 
length, which is often equal to that of the body of the beetle. Females 
with this type of genital tube would seem to be ill adapted for end-to- 
end mating with males because of the likelihood that the tube would be 
pulled out to its full length when mating individuals attempted to 
move in opposite directions. 

Oviposition. Oviposition has been observed for Lytta vesicatoria by 
Beauregard (1890), Xambeau (1900), and others and for cyanipennis 
by myself. It differs little from the oviposition process described for 
other Meloinae. In cyanipennis the burrow excavated by the female varies 
from one to two and one-half inches in depth. All burrows observed ran 
downward at about a 45 degree angle. On a few occasions females were 
observed to abandon partially completed burrows and, after a short 
period of feeding, to begin excavation of new ones. They seemed to 
excavate their burrows more frequently in darkness or subdued light 
than in bright light. 

Immediately after oviposition the female begins to pull soil down from 
the sides of the burrow to cover the egg mass. This activity continues 
for about ten minutes; at the end of this time the burrow is filled except 



16 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF LyttO 

for a slight depression marking the entrance. The female does not use 
the head to tamp down the soil, as has been reported for some species 
of the genus Epicanta (Horsfall, 1943). 

The only important difference between my observations of oviposition 
in cyanipennis and Beauregard's (1890) description of oviposition in 
vesicatoria is that the latter species "nearly always" places its eggs in two 
masses divided by a thin layer of soil. According to Beauregard, each of 
the masses is the product of a different ovary. The number of eggs pro- 
duced by a female of vesicatoria is given by Beauregard as 80 to 250. 
The number of eggs laid by cyanipennis is comparable. In both species 
females presumably oviposit only once, and they generally die within a 
few days after ovipositing. 

Development of Immature Stages 

Incubation. The average incubation period for eggs of the 15 species 
of Lytta for which data are available is 16 days. The shortest period 
recorded is 8 days (for cyanipennis) and the longest 27 days (for vesi- 
catoria; MacSwain, 1956 ) . Seasonal variation in the length of the incuba- 
tion period of vesicatoria was noted by Beauregard (1890). Eggs laid 
in June hatched in 21 days; eggs laid in July hatched in 17 to 18 days. 

Number of larval instars. Early students of the bionomics of the Me- 
loidae recognized four stages of larval development, not all of which 
correspond to separate instars. In recent times this procedure has been 
abandoned, but I have had to adopt it in the present discussion because 
of uncertainty as to the number of instars involved in larval develop- 
ment of Lytta. I am going to call the stages the first larval instar, grub 
stage, coarctate larval instar, and ultimate larval instar. In Beauregard's 
(1890) work on Lytta vesicatoria the first larval instar is termed the 
premiere larve or triongulin; the grub stage, the seconde or deuxieme 
larve; the coarctate larval instar, the pseudo-chrysalide; and the ultimate 
larval instar, the troiseme larve. 

Uncertainty as to the number of larval instars in Lytta stems from con- 
flicting reports of the grub stage or deuxieme larve. Lichtenstein (1879) 
definitely indicated that this stage is composed of only three instars 
(trois larves blanches), which would make the coarctate instar the fifth 
and the ultimate instar the sixth. The coarctate stage of the genus Lytta 
has been referred to several times as the fifth larval instar by subsequent 
workers, e.g., Linsley and MacSwain (1942), but presumably none of 
these workers has actually observed the complete development of a 
species of Lytta. 

In his account of rearings of vesicatoria, Beauregard (1890) men- 
tioned three molts between the first larval instar and the coarctate larval 
instar. Put another way, his account, like Lichtenstein's, indicates that 



BIONOMICS 17 

there are only three instars during the grub stage. However, in his treat- 
ment of the morphology of the larvae of vesicatoria, Beauregard (1890) 
described and figured four different larval forms in the grub stage. These 
are his seconde larve au troiseme jour, premiere mue de la seconde larve, 
deuxieme mue de la seconde larve, and troisieme mue de la seconde larve. 
On the basis of Beauregard's work these larval forms are sufficiently 
different morphologically that they almost certainly belong to separate 
instars. 

The discrepancy between the number of larval instars recorded by 
Beauregard in his account of rearings and the number described in his 
treatment of larval morphology has to be accounted for by a failure on 
his part to observe one of the larval molts during the grub stage. That 
he might indeed have done this is suggested by the following remarks 
given on page 293 of his account of rearings. 

Du 10 au 12, elle [the larva] subit d'assez profondes modifications; sa tete et 
surtout le premier anneau thoracique, grossissent beaucoup. La tete prend la 
position inclinee en bas, si caracteristique chez l'individu adulte, et les mandi- 
bules acquierent une coloration brune a la pointe. C'est, dans le developpement 
de cette larve, une phase que j'ai observee au cours de toutes mes educations, 
et qui marque un nouvel etat tres caracteristique de la seconde larve. 

In the few rearings of cyanipennis that I carried out several years ago, 
only three instars were observed in the grub stage. Yet, upon re-examin- 
ing my material I find striking morphological differences between a larval 
specimen recorded at the time of the rearings as an early fourth instar 
larva and one which was thought to be a late fourth instar larva. My 
findings are therefore like Beauregard's. Further, since the cast skins 
of larvae are relatively conspicuous in the grub stage, the only explana- 
tion I can offer for our apparent failure to record four instars in this 
stage is that the cast skin of one of the larvae is eaten by the larva of 
the following instar. 

While it is evident that no definite statement regarding the number 
of larval instars in the genus Lytta can be made, the weight of evidence 
suggests that the grub stage is composed of second, third, fourth, and 
fifth larval instars. On this basis the coarctate instar is the sixth, and the 
ultimate larval instar is the seventh. Careful studies of Epicauta (Hors- 
fall, 1941, 1942, and 1943) and of Mylahris (Paoli, 1938) have demon- 
strated the presence of seven definite larval instars in these genera. It 
would appear probable that this same number is also characteristic of 
the genus Lytta. 

Larval hosts. Bees recorded as larval hosts of Lytta are listed in Table 
I. Except for the species of Collet es ( Colletidae ) , all hosts are members 
of the family Anthophoridae. Data on the extent of parasitism by species 
of Lytta are available for a few nesting sites of bees. Linsley and Mac- 



18 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lytta 

Table I. Larval Hosts of Lytta 
Lytta species larval host authority 

L. (Paralytta) melaena Diadasia bituberculata (Cresson) Linsley & MacS wain (1952) 

L. {Paralytta) tenebrosa A nthophora linsleyi Timberlake Linsley & MacSwain (1942) 

L. (Paralytta) variabilis Ptilothrix sumichrasti (Cresson) Linsley, MacSwain, and 

Smith (1956) 

L. (Poreospasta) chloris A nthophora linsleyi Timberlake Linsley & MacSwain (1942) 

L. (Poreospasta) stygica Anthophora linsleyi Timberlake Linsley & MacSwain (1942) 

L. (Poreospasta) moerens Anthophora stanjordiani Cresson Linsley & MacSwain (1942) 

Colletes fulgidus Swenk MacSwain (1956) 

L. (Lytta) vesicatoria Colletes sp. Beauregard (1885) 



Swain (1942) found larvae of L. tenebrosa in 1.45 per cent, stygica in 
3.82 per cent, and chloris in 0.92 per cent of 759 cells of Anthophora 
linsleyi. They also reported L. melaena parasitizing 4.0 per cent of 377 
cells of Diadasia bituberculata (Linsley and MacSwain, 1952). Parasitism 
of Ptilothrix sumichrasti by L. variabilis was reported to be at a low 
(but unspecified) level at several nesting sites in Zacatecas, Mexico 
(Linsley, MacSwain, and Smith, 1956). 

The larva of Lytta chloris requires the contents of but a single cell of 
its host to complete development ( Linsley and MacSwain, 1942 ) . Larvae 
of tenebrosa, melaena, and vesicatoria probably need the contents of 
more than one cell each. Therefore, as pointed out by Linsley and Mac- 
Swain (1952), extent of parasitism determined by counting the number 
of bee cells containing larvae of a species of Lytta may not be an ac- 
curate indication of the effect of parasitism on the host bee. In all 
probability larvae of Lytta attack single cells when parasitizing bees of 
relatively large body size and attack two or three cells each when para- 
sitizing bees of smaller body size. In the former case they develop 
entirely on provisioned material. In the latter, they frequently encounter 
and attack immature stages of the bee. 

In the laboratory larvae of Lytta have been reared on the provisions 
of a variety of bees. Lichtenstein's ( 1875 ) first success in inducing larvae 
of vesicatoria to feed under laboratory conditions was achieved with 
honey from the esophagus of the honey bee (Apis mellifera Lin- 
naeus). Larvae were reared on this material to the late grub stage. 
Beauregard (1890) reared a larva of this same meloid to the early grub 
stage using Apis honey thickened by the addition of rose pollen. In 
other experiments larvae of vesicatoria were observed to feed on pro- 
visioned material of species of Andrena, Ceratina, Eucera, Halictus, 
Megachile, and Osmia ( Lichtenstein, 1875, 1877, and 1879; Beauregard, 
1890). Complete rearings were made with the Ceratina, Halictus, Mega- 
chile, and Osmia provisions. In my own rearings of cyanipennis (to the 



BIONOMICS 19 

coarctate larval instar) provisioned pollen and larvae and pupae of 
Norma melanderi Cockerell were used as food. 

First larval instar. Upon hatching, first instar larvae are pale in color. 
Within a few hours they darken and become very active, crawling rapidly 
over the ground and investigating holes and cracks in it. When touched 
they curl up into a ball. In vesicatoria they are said to be negatively 
phototactic (Beauregard, 1890). In cyanipennis, however, they are 
strongly positively phototactic; in the rearing cage they gathered in 
groups on the side nearest the source of light, and they would follow a 
light source at it was moved around the cage. As in the case of other 
meloids, first instar larvae of species of Lytta are capable of living an 
extended period of time while searching for nests of their hosts. Larvae 
of vesicatoria were kept alive at least one month without food by Beau- 
regard (1890), and larvae of cyanipennis that I studied were still healthy 
15 days after hatching (at which time they were given food). 

The means of entry of first instar larvae into the cells of their hosts 
has not been determined. It seems likely that entry is made before the 
bee has finished provisioning its cell and while the cell is still open. It 
has also not been determined whether the larva destroys the egg of the 
bee before attacking the pollen material. It has been shown, however, 
that it is not necessary for a larva to feed on an egg before eating pollen. 

Grub stage. Feeding of the first instar larva is completed in from one 
to ten days. The larval cuticle then splits along the epicranial suture and 
the line of dehiscence on the thorax, and the second instar larva emerges 
head first. With this instar the larva enters the grub stage. Larvae of 
cyanipennis in this stage observed by me fed alternately on pollen and 
the contents of the bodies of immature Nomia bees. While in the grub 
stage each of the meloid larvae consumed large amounts of pollen in 
addition to five to eight larval or pupal bees. From time to time they 
emitted long strings of yellow fecal material. 

When replete the grub stage larva becomes restless and soon begins 
excavation of its resting chamber. In the laboratory the recorded length 
of the grub stage is 14 to 19 days for vesicatoria (Beauregard, 1890) 
and 36 days for cyanipennis (personal observation). In both species the 
larva spends the last half of the stage lying inactive in the resting 
chamber. 

Coarctate larval instar. In the species of Lytta that have been studied, 
the coarctate larval instar is attained in late summer and normally lasts 
through the following winter. In the molt to this instar the cast skin 
of the grub stage larva is shed entirely except for a small portion that 
adheres to the tip of the abdomen of the coarctate larva. The coarctate 
larva is convex in form, with the appendages, as is usual for Meloidae 



20 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

in this stage, reduced to mere stubs. According to Lichtenstein (1879), 
coarctate larvae of vesicatoria normally exude from the abdominal seg- 
ments a clear liquid which subsequently hardens to form lenses, giving 
the body the appearance of being covered with diamonds. This phenom- 
enon has not been observed for other species of Lytta. 

Ultimate larval instar. The ultimate larval instar is reached in spring. 
Morphologically it is similar to the grub stage. In vesicatoria it lasts 
from 12 to 15 days, with an active period of 3 or 4 days (Beaure- 
gard, 1885 and 1890). In tenebrosa it lasts two or three weeks; during 
this time the larva constructs a pupal chamber near the surface of the 
soil (Linsley and MacSwain, 1942). Whether such a chamber is con- 
structed by the larvae of other species of Lytta is not known. 

Pupal stage. The pupal stage lasts an average of 15 days in vesicatoria, 
according to Beauregard (1890). Development of color in the pupa of 
vesicatoria was described in some detail by this same author (Beaure- 
gard, 1885). 



Systematics and Phylogeny 

Genus LYTTA Fabricius 

Cantharis, Geoffroy, 1762, Histoire abregee des insectes, vol. 1, p. 339. 
Oliver, 1795, Entomologie, ou histoire naturelle des insectes, Coleopteres, vol. 
3, no. 46, p. 1. Lacordaire, 1859, Histoire naturelle des insectes, genera des 
Coleopteres, vol. 5, p. 676. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, 
p. 103. Champion, 1892, Biol. Centrali-Americana, Coleoptera, vol. 4, pt. 2, 
p. 437. Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 293. Misapplication of 
Cantharis Linnaeus, 1758. 

Lytta Fabricius, 1775, Systema entomologiae . . . , p. 260. LeConte, 1853, 
Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 330. Escherich, 1895, Verhandl. 
Kaiserlich-Koniglichen Zool.-Bot. Ges., vol. 44 (1894), p. 251. MacSwain, 
1956, Univ. California Publ. Ent., vol. 12, p. 74. 

Pomphopoea LeConte, 1862, Smithsonian Misc. Coll., vol. 3, art. 136, p. 273. 
(See p. 222 of present work for additional references.) 

Poreospasta Horn, 1868, Trans. Amer. Ent. Soc, vol. 2, p. 139. 

Pubescence fine, never obscuring surface or contributing appreci- 
ably to over-all color of beetle. Upper surface of body sparsely pubescent 
to glabrous; under surface always pubescent. Antennae elongate, usually 
longer in male than in female, basically moniliform (modified to serrate 
in male of peninsula ris; subfiliform in female of peninsularis and in 
polita), not strongly compressed. Male antennae usually with intermedi- 
ate segments broader than in female, so that antennae are not as 
strongly clavate; in many New World species two or more of the inter- 
mediate segments incrassate (sometimes distorted also), as wide as or 
wider than distal segments. Female antennae moniliform clavate, the 
segments becoming more moniliform distally. Mouth parts unmodified 
mandibulate. Elytra and hind wings fully developed, except in sublaevis, 
which has shortened elytra and lacks hind wings. Mesepisterna meeting 
broadly on midline of body; marginal area ample. Under surface of 
thorax never with a red spot or suffusion at center. Legs frequently 
modified in male. Middle tibiae bowed laterad except in males of the 
Eucera and Mutilata groups of the subgenus Adicolytta. Outer hind 
tibial spur usually thicker than inner one, usually obliquely truncate. 
Tarsal claws cleft to base or (morrisoni and moesta) with ventral blade 
fused solidly to dorsal one basally and much shorter than it (as in the 
Eupomphini); blades smooth, never serrate. Male sixth abdominal ster- 
num emarginate, the emargination triangular or rounded. Male genitalia 
with gonoforceps uniformly sclerotized, without lightly sclerotized areas 

21 



22 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

at base of gonostyli; aedeagus primitively (and usually) with two well- 
developed ventral hooks, sometimes with only one, rarely with none. 

Type species. The type species of Lijtta is Meloe vesicatoria Linnaeus, 
as fixed by subsequent designation by Wellman (1910, p. 392). Accord- 
ing to Wellman, "vesicatoria has by several authors (v. Escherich, Ver. 
k. k. zool.-bot. Gesell, 1894, p. 19) been designated as the type of the 
genus Lytta F." However, I have been unable to find any designation 
before Wellman's. 

No type designation is needed for Cantharis of Geoffroy and subse- 
quent authors who applied this name to species of Meloidae. Its usage 
in this manner is nothing more than a misapplication of the name. 

Pomphopoea and Poreospasta are herein regarded as subgenera of the 
genus Lytta, and their type species are indicated in appropriate sections 
of the systematic treatment that follows. 

Remarks. The generic synonymy given above is complete in that all 
synonyms are included. It is incomplete in that only those references of 
major significance are cited. A more nearly complete list of references 
was given by Borchmann ( 1917 ) . The only major work not cited in his 
synonymy is MacSwain's (1956) study of the first instar larvae of Lytta. 

Limits of the Genus 

The genus Lytta is a member of the tribe Lyttini of the subfamily 
Meloinae (or Lyttinae, if Meloe is placed in a separate subfamily). This 
tribe includes, in addition to Lytta, some 26 named genera as well as 
several other genera still undescribed. Taxonomically the tribe is so 
poorly understood that it is quite impracticable to speculate on the phy- 
logenetic relationships of the genera within it. In the past it has served 
as a convenient lumping-place for Meloidae not assignable elsewhere. 
Relatively recently attention has been called to its polyphyletic nature, 
two groups of species having been removed from the tribe following a 
study of their first instar larvae. Further restriction of the Lyttini is sug- 
gested in the following discussion. 

One of the most vexing problems encountered in the course of this 
study has been that of establishing the limits of the genus Lytta. If the 
fauna of a single country or district is considered, it is relatively easy to 
draw up a diagnosis of the genus and to place it quite satisfactorily in a 
key to genera. As the geographic scope of treatment is increased, it 
becomes more and more difficult to characterize the genus and to 
separate it from others. Like others, I have been unable to find any single 
character or simple combination of characters that will distinguish mem- 
bers of the genus from other Meloidae, and I have therefore been forced 
to use a lengthy list of characters in framing my diagnosis of the genus 
( above ) . 



SYSTEMATICS AND PHYLOGENY 23 

In North America, the genus Li/tta as herein defined has essentially 
the same composition as it does in current catalogs ( Blackwelder, 1939 
and 1945) except for: 1) inclusion of moesta and morrisoni, previously 
transferred to the genus from the Eupomphini (Selander, 1954); 2) re- 
moval by MacSwain ( 1951 ) of sphaericollis ( Say ) , compressicornis 
(Horn), convexa (LeConte), and suavissima (Wellman) to form the 
genus Linsleya MacSwain in the tribe Epicautini; 3) inclusion of Poreo- 
spasta sublaevis (= polita), as first suggested by MacSwain (1956); 
4) inclusion of the species of Pomphopoea; and 5) removal of the 
Central American sanguineo guttata Haag-Rutenberg to form the mono- 
typic genus Paniculolytta Selander ( 1960 ) . 

The epicautine affinities of Linsleya have been satisfactorily demon- 
strated by MacSwain. The fact that the true relationships of the genus 
were not suspected until MacSwain's study of the first instar larva of one 
of the species is indicative of the difficulty of the problem of establishing 
generic limits and relationships within the tribe Lyttini on the basis of 
adult morphology alone. 

MacSwain ( 1956 ) has also shown the close relationship existing be- 
tween sublaevis and the group of western North American species of 
Lytta associated with stygica. I am in complete agreement with his 
decision to place Poreospasta in Lytta, but I feel that sublaevis and its 
relatives are sufficiently distinct that Poreospasta merits subgeneric rank. 

Pomphopoea has almost invariably been accorded full generic status, 
and if I were to consider only the New World species of Lyttini, I would 
undoubtedly continue to regard it as a valid genus. However, its species 
are apparently more closely related to Lytta vesicatoria, the type of 
Lytta, than are any of the North American species of Lytta. This is 
confirmed by adult and larval morphology and by the similarity of the 
feeding habits of adults of vesicatoria and Pomphopoea. It is there- 
fore necessary either to follow the arrangement suggested in the present 
work and reduce Pomphopoea to subgeneric rank or, less desirably, to 
elevate the other subgenera of Lytta to full generic rank. 

Removal of sanguineo guttata from Lytta is arbitrary. The species 
agrees with the members of the subgenera Paralytta and Adicolytta in 
having incrassate male intermediate antennal segments, but there is no 
other basis for assuming that the species is more closely related to Lytta 
than to a number of other lyttine genera. 

With the reduction of Poreospasta and Pomphopoea to subgenera of 
Lytta and removal of the genus Pyrota Dejean from the Lyttini (Mac- 
Swain, 1956), only two genera of Lyttini are now represented in North 
America: Lytta and Paniculolytta. 

Lytta differs from Paniculolytta in the following characters: form much 
less elongate; male intermediate antennal segments not greatly length- 



24 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

ened; male trochanters lacking a heavy tuft of stiff setae; aedeagus 
almost always with one or two ventral hooks. Some species of Lytta 
superficially resemble species of Pyroto, but the latter may be distin- 
guished by their slender, tapered, often somewhat geniculate antennae, 
by the separation of the mesepisterna of the thorax, and by the fact 
that the posterior margin of the male sixth abdominal sternum is fre- 
quently turned sharply dorsad and is membranous. In addition, males 
of most species of Pyrota have the maxillary palpi (particularly the last 
segment) enlarged and modified in form. Lytta sublaevis, the only ap- 
terous species of the genus, is similar in general appearance to members 
of the genus Meloe. It is readily distinguished from them by its orange 
frontal spot, exposed scutellum, and in many cases the presence of red or 
orange on the vertex or pronotum or both. 

For South America, Denier (1940) listed 12 species in Lytta, although 
he did not regard any of them as actually congeneric with the type of the 
genus. Subsequent work has confirmed this conclusion. In 1959, Kaszab 
removed the species binotatithorax Pic ( = seminigra Borchmann ) , 
bruchi Pic, catamarcensis (Brethes), colon ( Burmeister ) , dimidiata Fis- 
cher ( = muelleri Borchmann ) , griseopubescens Denier, nigropicta 
Denier, and quadrilineata ( Haag-Rutenberg ) ( = bilineatipennis Pic ) 
from Lytta and placed them in a new genus, Acrolytta Kaszab. My studies 
confirm Kaszab's and, in addition, indicate that the species abbreviata 
Klug, inflaticeps Beauregard (= fissiceps Haag-Rutenberg and adonis 
Pic), and philippi (Reed) should be removed from Lytta. New genera 
are proposed for these species elsewhere (Selander, 1960). With these 
changes the only South American species of Meloidae still assigned to 
Lytta is neivai Denier. I have not seen specimens of this species, but on 
the basis of its description, it will probably have to be removed from 
the genus also. 

The relationships of the South American lyttines are obscure. Kaszab 
regards Acrolytta as most closely related to the Ethiopian genus Afrolytta 
Kaszab, but I fail to see the basis for this conclusion. All South American 
species of Ly ttini that I have examined ( including species of Pseudomeloe 
Fairmaire and Germain and Picnoseus Solier) agree in having a lightly 
sclerotized or membranous area on each of the gonostyli of the male 
genitalia. This fact leads me to believe that the lyttine fauna of South 
America may have been derived from a single ancestral type. In any 
event, I have found no evidence of close relationship between the genus 
Lytta and any of the South American lyttines. 

In the Old World Lytta presently contains a very heterogeneous assem- 
blage of species and is badly in need of critical revision. I have not 
studied enough material to justify making extensive formal changes in 
the classification of the Old World representatives of the genus. 



SYSTEMATICS AND PHYLOGENY 25 

Of the 23 species of Meloidae of the Ethiopian region formerly assigned 
to the genus Lytta, 7 were recently transferred from the genus by Kaszab 
( 1959 ) . The species lucida Haag-Rutenberg, nitidula Fabricius, pallidi- 
pennis Haag-Rutenberg, semilineata Haag-Rutenberg, and sumptuosa 
(Castelnau) are placed by Kaszab in the genus Prolytta Kaszab, while 
the species amoena (Peringuey) and mashuna (Peringuey) form the 
genus Afrolytta Kaszab. These transfers are supported by my studies of 
several of the species involved, although I do not share Kaszab 's opinion 
that Prolytta is a lyttine genus. Rather, it seems necessary to erect a new 
tribe, Prolyttini, for this genus on the basis of the following combination 
of characters: wings with vein 2ndA 2 present; sixth abdominal sternum 
hardly visible behind fifth; male gonostyli fused to apex; female genitalia 
lacking gonostyli. 

If the characters of the species spilotella (Peringuey) are at all repre- 
sentative of the Ethiopian Spilotella Group as a whole, then it seems 
highly probable that this group should also be removed from the genus 
Lytta and from the tribe Lyttini. In addition to the nominate species, 
this group contains the species elegantula (Peringuey), enona (Perin- 
guey), namagua Kaszab, rubrolineata Kaszab, szekessyi Kaszab, and 
vellicata Erichson. Its affinities would seem to be more epicautine than 
lyttine. 

Few if any of the other Ethiopian species placed in Lytta are likely 
to be retained in this genus when they have been critically studied. These 
species are bayoni Pic, benguellana Pic, brucci Castelnau, cantharoides 
Thunberg, carneola (Peringuey), exclamans (Fairmaire), fryi (Wollas- 
ton), myrmido (Fairmaire), and validicornis (Fairmaire). 

The Eurasian fauna includes a total of 43 species of Meloidae assigned 
to the genus Lytta at the present time. The only one of these found in 
Europe is vesicatoria, which also has an extensive range in Asia. Six of 
the species (apicalis, aeneiventris, spissicomis, fissicollis, nigrifinis, and 
usta) occur in the Oriental region; the rest are Palearctic in distribution. 
The relationships of 20 of the species are sufficiently clear, either from 
data given in the literature or from my personal examination of speci- 
mens, that they may be assigned to subgenera within Lytta. These species 
fall into five subgenera, four of which seem to be confined in distribution 
to Asia (see discussion in the following sections). The relationships 
of the remaining 23 species are unknown to me. Many of them are 
represented only by type material which has not been examined since 
the original descriptions were made, and it seems likely that some are 
synonyms of better-known species. A number of them probably will be 
transferred to other genera eventually; others are probably assignable 
either to the subgenera defined in the present work or to new subgenera. 
Clarification of the systematic position of these species will be a major 



26 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lljttd 

step in the advancement of our knowledge of the generic classification of 
the Lyttini. The species in question are badakschanica Kaszab, hieti 
Wellman, cardonii ( Fairmaire ) , dichroa Fischer von Waldheim, disci- 
pennis (Fairmaire), fissicollis (Fairmaire), flava Thunberg, flavicollis 
Gyllenhal, flavoangulata (Fairmaire), flaviventris Ballion, fulviceps 
(Maklin), ghbricollis ( Waterhouse ) , impressicollis Fairmaire, laeta 
( Waterhouse ) , malatiensis (Heyden), manicata Sahlberg, marginicollis 
Haag-Rutenberg, melanura Hope, nigrifinis (Walker), punctata Pallas, 
rubra Hope, thibetana Olivier, and usta (Fairmaire). 

Classification 

Conspectus. The genus Lytta is herein divided into 8 subgenera, 18 
species groups, and 11 species subgroups. The proposed arrangement of 
these taxa is as follows. 

Subgenus PARALYTTA, new subgenus 
Nitidicollis Group 
Fulvipennis Group 
Variabilis Group 
Variabilis Subgroup 
Koltzei Subgroup 
Quadrimaculata Subgroup 
Reticulata Group 
Reticulata Subgroup 
Deserticola Subgroup 
M agister Group 

Tenebrosa Subgroup 
Melaena Subgroup 
Magister Subgroup 
Subgenus ADICOLYTTA, new subgenus 
Peninsularis Group 
Eucera Group 
Mutilata Group 
Subgenus POREOSPASTA Horn, new status 
Stygica Group 
Moerens Group 
Moerens Subgroup 
Cyanipennis Subgroup 
Childi Subgroup 
Luteovittata Group 
Fairmairei Group 
Clematidis Group 
Subgenus POMPHOPOEA LeConte, new status 
Polita Group 



SYSTEMATICS AND PHYLOGENY 27 

Aenea Group 
Sayi Group 

Subgenus LYTTA Fabricius 
Vesicatoria Group 
Menetriesi Group 

Subgenus PSEUDOLYTTA, new subgenus 

Subgenus MESOLYTTA, new subgenus 

Subgenus INDIOLYTTA, new subgenus 

Basis. The present classification is based primarily on morphological 
characters of the adult beetles. However, data from morphology of lyttine 
larvae in the first instar given by MacSwain (1956) have been of great 
value in its formulation. First instar larvae have been described for 8 
species of Paralytta, 2 of Adicolytta, 14 of Poreospasta, 2 of Pomphopoea, 
and 1 of Lytta (MacSwain, 1956; Selander, 1959). At the generic-sub- 
generic level the present classification differs from MacSwain's in that 
Pomphopoea is not accorded generic status. Yet MacSwain did call atten- 
tion to the close relationship of Pomphopoea and Lytta indicated by his 
larval studies. It seems probable that all the subgenera of Lytta can be 
defined in the larval stage, although rather lengthy combinations of 
characters will be required for most of them. MacSwain's work contains 
a diagnosis of Pomphopoea and I have given a tentative diagnosis of 
Adicolytta (Selander, 1959). 

Old World subgenera. The subgenera Lytta, Pseudolytta, Mesolytta, 
and hidiolytta are confined to the Old World and are not treated in 
detail in the present work. Their diagnostic characters are given in the 
key to subgenera that follows this discussion. Some additional informa- 
tion concerning them is given below. 

The subgenus Lytta is divided into two groups. The Vesicatoria Group 
is characterized by having two spurs on each of the male middle tibiae 
and by the presence of mesal hooks on the gonostyli of the male genitalia. 
This group includes the species vesicatoria (Linnaeus), roborowski 
Dokhtouroff, suturella Motschulsky, caraganae Pallas, and sifanica Se- 
menov. The Menetriesi Group has only a single spur on each of the 
male middle tibiae and lacks mesal hooks on the male gonostyli. The 
species of this group are menetriesi Falderman, flavipennis Motschulsky, 
antennalis (Marseul), and regis-zahiri Kaszab. The species of the sub- 
genus form a very closely knit taxonomic unit. They were last treated by 
Escherich (1895). 

The subgenus Pseudolytta, new subgenus, corresponds exactly to the 
Aeneiventris Group of Maran (1941). Its type species is hereby desig- 
nated as Lytta aeneiventris Haag-Rutenberg. Other species included in 
the subgenus are kwanhsiensis Maran, taliana Pic, and spissicornis (Fair- 
maire ) . 



28 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

The subgenus Mesolytta, new subgenus, is erected for two species of 
uncertain relationships. These are Lytta coccinea Menetries, which is 
hereby designated as type species, and deserti Semenov. 

The subgenus Indiolytta, new subgenus, contains a single species, 
Lytta apicalis Haag-Rutenberg, which automatically becomes its type 
species. 

Key to Subgenera x 

1. Labrum deeply emarginate (Figs. 155-58); fore femora concave on 
anterior side; male middle tibiae each with posterior spur long, 
curving strongly posteriad; gonostyli of male genitalia (Figs. 348-50) 
slender, glabrous, lacking mesal hooks; male intermediate antennal 
segments not incrassate; metallic green, blue, or aeneous beetles with 

femora and at least hind tibiae orange. North America 

Pomphopoea ( p. 222 ) 

Labrum not deeply emarginate; fore femora not concave on anterior 
side; male middle tibiae with posterior spurs not modified as above; 
gonostyli of male genitalia variable; male intermediate antennal seg- 
ments incrassate or not; if body metallic in color, femora and tibiae 
dark colored 2 

2. Male intermediate antennal segments incrassate, with IV and V as 
wide as or wider than distal segments, or at least with IV wider than 
V; male fore tibiae each with two spurs; male hind trochanters neither 
spined nor angulate; gonostyli of male genitalia glabrous or essentially 
so, lacking mesal hooks; aedeagus generally with two well-developed 

ventral hooks, rarely with only one. North America 3 

If male intermediate antennal segments incrassate (some North 
American species of Poreospasta) , then male hind trochanters angu- 
late or spined or gonostyli of male genitalia clothed with long setae, 
or both 4 

3. Male intermediate antennal segments somewhat compressed and usu- 
ally flattened on inner side but never asymmetrically produced or 
excavate; male middle tibiae sometimes impressed subapically and 
first segment of male middle tarsi sometimes weakly distorted, but 
male legs otherwise unmodified; gonostyli of male genitalia variable 

1 This key and the keys to groups, subgroups, and species given in the sections 
on the North American subgenera are designed to summarize the principal differences 
between taxa, without regard to ease of determination of individual specimens. Males 
should run satisfactorily in them; females cannot be run to subgenera (except for 
Pomphopoea). For the determination of females and as a convenience in the deter- 
mination of males of the North American species of Lytta, an artificial key is pro- 
vided, beginning on page 35. 



SYSTEMATICS AND PHYLOGENY 29 1 

but distinct in form from those of Adicolytta; male head often modi- 
fied behind eyes. Southwestern Canada to Central America 

Parolytta (p. 49) 

At least segments IV and V of male antennae conspicuously, asym- 
metrically distorted, except in the southern Mexican erebea; male 
trochanters with modified pubescence and male middle tibiae and 
either fore tibiae or middle femora modified for holding female dur- 
ing courtship, except in the Baja Californian peninsularis; gonostyli 
of male genitalia slender, either evenly rounded at apex or weakly 
lobed before apex (Figs. 324-30); male head never modified behind 
eyes. Primarily Mexican-Central American in distribution, with muti- 

lata extending north into the southwestern United States 

Adicolytta (p. 137) 

4. Male fore tibiae each with two spurs, or if with only a single spur 
(two North American species), the hind trochanters spined. North 

America and Asia 5 

Male fore tibiae each with only a single spur; male hind trochanters 
not modified. Eurasia 6 

5. Gonostyli of male genitalia each bearing a mesal hook, or if hooks 
absent, then gonostyli clothed ventrally with long setae. North 

America and Asia Poreospasta (p. 157) 

Gonostyli of male genitalia lacking mesal hooks, glabrous; female 
pygidium produced. India Indiolytta (p. 28) 

6. Male middle tibiae each lacking posterior (inner) spur; male head 
deeply grooved behind eyes; head dark without a pale frontal spot; 
colors not metallic; antennal segments distinctly beadlike, well sepa- 
rated; first and second segments of male tarsi strongly distorted, 
modified for clasping; gonostyli of male genitalia entirely glabrous, 

lacking mesal hooks. China Pseudolytta (p. 27) 

Male middle tibiae each with two spurs or lacking anterior (outer) 
spur; male head not grooved; head, if dark, usually with a pale 
frontal spot; second segment of male fore tarsi not modified; gono- 
styli of male genitalia sparsely clothed with setae (setae very short 
in Mesolytta ) 7 

7. Colors not metallic; head, if dark, lacking a pale frontal spot; gono- 
styli of male genitalia clothed with veiy short setae, lacking mesal 

hooks. Southwestern Asia Mesolytta (p. 28) 

Head and body metallic green or blue; head with a pale frontal spot; 
gonostyli of male genitalia clothed ventrally with long setae, with or 
without mesal hooks. Eurasia Lytta (p. 27) 



30 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 



Eurasia 



Asia 



N.A. 



N.A. 



Asia 
+ N.A. 



Asia 



Asia 



Fig. 1. Phylogeny of the subgenera of Lytta. 



Phylogeny of the Subgenera 

The suggested phylogenetic relationships of the subgenera of Lytta 
are shown in Figure 1. Two main lines may be recognized, one in North 
America which gave rise to Paralytta and Adicolytta, and another, prob- 
ably originating in Asia, which produced Poreospasta, Pomphopoea, 
Lytta, and Pseudolytta. The relationships of Mesolytta and Indiolytta are 
extremely doubtful. These subgenera may have evolved as early branches 
of the second main line or they may represent offshoots from the Lytta 
stock which formed before the divergence of the two main lines. The 
first main line is characterized by having the male intermediate antennal 
segments at least weakly incrassate and the male genitalia simple in form 
and glabrous. Inasmuch as both Paralytta and Adicolytta feed to a great 
extent on Convolvulaceae, preference for or specificity to this family of 
plants was probably characteristic of their common ancestor. In Para- 
lytta there has been divergence in different groups to such families as 
Compositae, Labiatae, and Papaveraceae. In Adicolytta one species has 
moved to Cucurbitaceae. 

The second main evolutionary line of the genus lacks the antennal 
modification of the first except in a few North American species of 



SYSTEMATICS AND PHYLOGENY 31 

Foreospasta in which the condition is clearly due to convergence with the 
first main line rather than to relationship. Close relationship between 
the subgenera Foreospasta and Lytta is indicated by the similarity of 
the male genitalia in the more primitive members of each (the gonostyli 
have mesal hooks and are pubescent). In the adult stage Pomphopoea has 
the general facies and color of the subgenus Lytta as well as antennal 
similarities; in the larval stage, according to MacSwain (1956), there 
are several distinctive characters shared by the two subgenera. It is 
therefore logical to suppose that the two are closely related. Pseudolytta 
is similar to the subgenus Lytta in general facies also and in addition 
shows modifications of the male legs reminiscent of those in the latter 
subgenus. 

On the basis of the phylogeny postulated in Figure 1, the earliest mem- 
bers of the second main line of the genus Lytta probably fed on Le- 
guminosae. In Poreospasta the Moerens Group is apparently restricted 
to this family, while the Stygica Group has developed the ability to feed 
on Compositae and Papaveraceae also. Both Pomphopoea and the 
subgenus Lytta feed principally on shrubs and trees. In the subgenus 
Lytta recorded host families are Leguminosae, Oleaceae, Salicaceae, and 
Caprifoliaceae. Potnphopoea feeds mainly on Rosaceae. 

Distribution 2 

The genus Lytta is primarily temperate in distribution, with only a few 
tropical representatives. Like other genera of Meloinae, it is not well 
represented on islands and does not occur at all on oceanic ones. 

In the Old World the genus ranges through most of Eurasia, from 
Spain and England to Manchuria, Japan, southeastern China, and Ceylon. 
The subgenus Lytta is very widely distributed. Mesolytta occurs in the 
Transcaspian region; Indiolytta, in India; Poreospasta, in southwestern 
Asia; and Pseudolytta, in eastern China. 

In the New World the genus is apparently limited to the North Ameri- 
can continent, where it ranges from Canada to Panama. The distribution 
in North America is considered in detail in the systematic sections that 
follow. Consequently, the present discussion is limited to a summary of 
the main distributional features of the subgenera and groups. Reference 
will be made to the maps shown in Figures 2-9. These were prepared by 
superimposing the ranges of the species for the various subgenera or 
groups. They are designed to show the total known range of each of 
the taxa as well as to indicate the pattern of distribution of the component 
species within this range. 

Most of the North American species of Lytta occur in the arid and 

- South American and Ethiopian species currently assigned to Lytta have been 
excluded from consideration in this section; see discussion of limits of the genus. 



32 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

semiarid regions of the southwestern United States and the Central 
Plateau of Mexico. Two main centers of distribution are evident: one in 
California (32 of the 51 species recorded from the United States occur 
there) and the other at the southern end of the Central Plateau. 

The subgenera Paralytta and Adicolytta probably diverged from a 
common ancestor in North America. In Paralytta the Variabilis Group 
(Fig. 2) has undergone most of its evolution in southern Mexico. The 
Magister Group (Fig. 3) presumably developed in the lowlands of 
California and only later entered the Sierra Nevada, the Sonoran Desert, 
and the Great Basin. The Reticulata Group (Fig. 4) is an offshoot of 
the Variabilis Group which apparently evolved in the more northern area 
it now occupies. The Fulvipennis and Nitidicollis groups (Fig. 5) are 
probably relicts of early, rather unsuccessful lines; their precise relation- 
ships are not clear. 

Adicolytta (Fig. 6) probably originated in southern Mexico. Its least- 
derived species (peninsidaris) is found at the southern tip of the penin- 
sula of Baja California, where it is obviously relict. On the Mexican 
mainland the least-derived form is erebea, which is known only from 
the mountains in the state of Mexico. This species is most closely re- 
lated to eucera, which ranges from northern Mexico to Panama. Eucera 
is the only species of the subgenus known to have deviated from the 
Convolvulaceae feeding habit, and its success as a species compared 
with that of erebea may be connected with this fact. Mutilata is remark- 
able as the only Mexican species of Lytta that crosses the Sonoran Desert 
and extends to the West Coast. 

Poreospasta probably reached North America as an immigrant from 
Asia, where it is now represented by three species groups. In North 
America the subgenus is represented by two large groups. According 
to my phylogenetic conclusions, the Moerens Group ( Fig. 7 ) has its more 
primitive members confined to the Pacific Coast. Three wide-ranging 
Rocky Mountain forms (cyanipennis, viridana, and nuttalli) assigned to 
the group would probably be placed in a separate group were it not for 
the fact that another species (nigrocyanea) is an almost perfect con- 
necting link between them and the Pacific Coast forms. The species 
nuttalli is noteworthy in extending farther east than any species of Lytta 
outside the subgenus Pomphopoea and in having a disjunct population 
in the Sierra Nevada of California. An interesting point here is that 
while nuttalli occurs with cyanipennis in Idaho, Utah, and in the Sierra 
Nevada, it apparently cannot follow it through Oregon and Washington. 

Except for the lack of a Rocky Mountain element and an extension by 
two species into the Sonoran Desert, the Stygica Group (Fig. 8) has a 
range much like that of the Moerens Group. All the species of the Stygica 
Group occur in California, and only three extend beyond the limits of 



SYSTEMATICS AND PHYLOGENY 



33 




>7 


'S »-v 


\ "* 


%A— 


<& . 


^ '^ 


yii ^ \ 






MAGISTER GROUP 






3 








NI T !OICOLLIS (left) 
8 FULVIPENNIS (right) 
GROUPS 



Figs 2-5. Distribution of the groups of the subgenus Paralytta. 



34 



BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lyttd 




Fig. 6. Distribution of the subgenus Adicolytta. 

Figs 7-8. Distribution of groups of the subgenus Poreospasta in North America. 

Fig. 9. Distribution of the subgenus Pomphopoea. 



SYSTEMATICS AND PHYLOGENY 35 

that state. Most of the species have a small range, and in general they 
are quite similar morphologically and ecologically. In southern Cali- 
fornia as many as seven species may occur at a single locality. Just what 
mechanism is involved in maintaining species limits within the group is 
perhaps the most intriguing unsolved problem in the genus Lytta. 

Pomphopoea (Fig. 9) is a very distinctive element in the North Ameri- 
can fauna, both morphologically and geographically. It evidently evolved 
from an immigrant stock which reached the North American Continent 
from Asia at a time when the deciduous forest was continuous through 
the Holarctic region. It is now restricted to the eastern United States, 
where it replaces the other subgenera of Lytta. Details of distribution of 
aenea and perhaps also of sayi would seem to indicate that at one time, 
probably following Pleistocene glaciation, there was a western expansion 
of range of the subgenus which has since been followed by a contraction, 
leaving isolated populations to the west of the present continuous range. 

In North America important barriers to dispersal of the genus Lytta 
are extremely arid conditions such as those of the Sonoran and Mojave 
deserts, large mountain ranges, and tropical lowlands. The West Coast 
of the United States has been very effectively isolated. In a few groups 
inhabiting this area species have entered the Sierra Nevada, but only 
one line (the Cyanipennis Subgroup of Poreospasta) has succeeded in 
entering the Rocky Mountains, while only one species (mdnerata) has 
penetrated into the Great Basin. The Great Plains have been invaded by 
only two species (reticulata and biguttata). The eastern United States 
shows no western or Mexican influence whatsoever. The general distri- 
butional picture of Lytta in North America, then, is one of peripheral 
centers from which movement has been rather limited, so that the center 
of the continent is depauperate in species. 

It is interesting to note that the distributional patterns displayed by 
Lytta in North America have little in common with those of some of the 
other genera of North American Meloidae. For example, in Nemognatha, 
which was recently revised by Enns (1956), the central area of the 
United States has a fairly good representation of species, only one small 
group is centered in California, and no group shows a definite southern 
Mexican center of distribution. Furthermore, except for one southeastern 
species (which is probably of Antillean origin), the eastern fauna is 
closely related to the western, apparently without close Old World affini- 
ties. 

Artificial Key to the North American Specks of Lytta 

1. Wingless, with shortened elytra. California sublaevis (p. 163) 

Wings and elytra fully developed 2 



36 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

2. Bright metallic green, blue, aeneous, or brassy beetles with femora 
and at least hind tibiae orange; labrum deeply emarginate (Figs. 

155-58 ) . Eastern North America 3 

If body metallic in color, femora and tibiae metallic or black; la- 
brum not or only shallowly emarginate 6 

3. Elytra dull brassy; fore and middle tibiae black; male fore tibiae 
each with a broad impression on anterior side and with only a single 

spur. Southeastern United States polita (p. 226) 

Elytra green, blue, or aeneous; all tibiae orange, at least at middle; 
fore tibiae lacking impression, each with two spurs 4 

4. Male fore tibiae thickened, stouter than middle tibiae, bowed; 
female sixth abdominal sternum deeply grooved medianly at apex 

(Figs. 288-89) 5 

Male fore tibiae not thickened, no stouter than middle tibiae, not 
bowed; female sixth abdominal sternum not or only feebly grooved 
aenea ( p. 229 ) 

5. Tibiae entirely orange; labral emargination extending to middle 
( Fig. 158 ) ; male fore femora contorted. Southeastern United States 

unguicularis ( p. 238 ) 

Tibiae narrowly black at base and apex; labral emargination not 
extending to middle (Fig. 157); male fore femora not contorted. 
Northeastern United States sayi (p. 235) 

6. Elytra strongly reticulate, with network of raised lines and enclosed 
cells clearly visible without magnification; elytra (except, rarely, 
extreme base) and under surface black; femora sometimes orange 

( with black base and apex ) 7 

Elytra rugose, appearing granular or smooth without magnification; 
if elytra and under surface black, femora never orange 17 

7. Tarsal pads (pale pubescence on ventral surface of tarsi) com- 
pletely absent; head, pronotum, femora, and tibiae entirely orange; 

rest black. Southwestern United States and northern Sonora 

magister ( p. 128 ) 

Tarsal pads present; color pattern not as above 8 

8. Pronotum hexagonal, strongly angulate at sides 9 

Pronotum circular or oval, rounded at sides 11 

9. Pronotum spinose on lateral margin of disk on each side 10 

Pronotum not spinose vulnerata (p. 130) 

10. Head and pronotum orange; emargination of male sixth abdominal 
sternum deep, rounded (Fig. 248); a rare variant. Western United 
States east of the Sierra Nevada vulnerata (p. 130) 



SYSTEMATICS AND PHYLOGENY 



37 



Head and pronotum red; emargination of male sixth abdominal 
sternum shallow, very obtusely angulate at apex. Isla Santa Marga- 
rita, Baja California Sur margarita (p. 136) 

11. Femora orange with black base and apex 12 

Femora black 13 

12. Elytra coarsely reticulate, most of the cells at least .5 mm. in diam- 
eter; head with orange confined to dorsal margin (including tem- 

pora) and a large frontal spot. San Luis Potosi and Hildalgo 

corallifera (p. 112 ) 

Elytra finely reticulate, most of the cells less than .5 mm. in diam- 
eter; head orange with black at sides of frontal area. Southwestern 
United States and Sonora deserticola (p. Ill ) 

13. Pronotoum black at center, broadly orange at sides; head with a 
large frontal spot. Southern Texas to Durango. . . .cribrata (p. 109) 
Color pattern not as above 14 

14. Pronotum black. Great Plains south to the Rio Grande 

reticulata ( p. 103 ) 

Pronotum orange 15 

15. Head, orange. Northeastern Arizona agrestis ( p. 105 ) 

Head black 16 

16. Pronotum subcircular (Fig. 175); elytra coarsely reticulate, most 
of the cells at least .5 mm. in diameter. Rio Grande region, Texas 
mirifica ( p. 107 ) 

Pronotum transversely oval (Fig. 176); elytra finely reticulate, most 
of the cells less than .5 mm. in diameter. Northwestern Arizona. . . 
arizonica ( p. 108 ) 

17. Ventral blade of each tarsal claw fused basally with dorsal blade 
and much shorter than it, appearing as at ventral tooth. California 

18 

Tarsal claws cleft to base, with blades free, subequal in length ... 19 

18. Elytra moderately densely clothed with semierect setae; tarsal claws 

distinctly curved; pronotum often orange morrisoni (p. 55) 

Elytra essentially glabrous; tarsal claws nearly straight; pronotum 
always black moesta (p. 56) 

19. Elytra partially or entirely red, orange, or yellow 20 

Elytra entirely black or metallic 31 

20. Elytra entirely pale in color 21 

Elytra partially black or dark brown 27 

21. Under surface and legs largely or entirely orange 22 

Under surface and legs entirely black. . 24 



38 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Luttd 

22. Body and legs entirely orange; length, 16 mm. -27 mm. Michoacan 

Morelos, and Guerrero sanguined (p. 148) 

At least tarsi black or brown; pronotum usually with a pair of black 
spots on disk 23 

23. Head black or largely so; male pygidium strongly, narrowly pro- 
duced at middle ( Fig. 209 ) ; an uncommon variant. Southern Mexico 

quadrimaculata ( p. 93 ) 

Head orange; male pygidium not so produced. Arizona, New 
Mexico, Sonora, and Chihuahua biguttata (p. 86) 

24. Head and pronotum black 25 

Head and pronotum orange 26 

25. Elytra a rich dark red; head and pronotum sparsely punctate and 

pubescent. Southern Mexico cardinalis (p. 155) 

Elytra orange; head and pronotum densely punctate and pubescent. 
Nebraska to Texas and (probably) Coahuila. . .julviyennvi (p. 58) 

26. Pronotum transverse, with a pair of black spots on disk at middle. 

Texas and northern Tamaulipas lecontei (p. 60 ) 

Pronotum elongate, immaculate. Southern California and northern 
Baja California Norte nitidicollis (p. 54) 

27. Elytra each with a broad brown discal vitta which begins at or near 
humerus and extends to or near to apex. Sonoran Desert region of 

Sonora sonorae ( p. 85 ) 

Elytra not as above 28 

28. Elytra black, each with lateral margin and a broad discal vitta 
yellow (Fig. 161). Puebla, Guerrero, and Oaxaca. . .scitula (p. 90) 
Elytra not as above 29 

29. Elytra black at apex. Southern Mexico proteus (p. 97 ) 

Elytra orange or yellow at apex 30 

30. Elytra (Fig. 159) each usually with a black spot at apical third and 
another at basal third, rarely with basal spot missing; elytral suture 
pale (except where spots may reach it); male pygidium strongly, 
narrowly produced; female sixth abdominal sternum strongly 
cupped. Southern Chihuahua and the Rio Grande to southern 

Mexico quadrimaculata (p. 93 ) 

Elytra (Fig. 160) never with a black spot at basal third; elytral 
suture usually black for basal half; male pygidium not produced 
as above; female sixth abdominal sternum not strongly cupped .... 
biguttata (p. 86) 

31. Pronotum partially or entirely yellow, orange, or red 32 

Pronotum entirely black or metallic 51 



SYSTEMATICS AND PHYLOGENY 39 

32. Pronotum strongly angulate at sides (Fig. 184), orange with a 
black or piceous anterior marginal area which sometimes extends 
posteriad as a median stripe; vertex orange, almost always with a 
median black stripe or fine line; rest of body and legs black, with- 
out metallic luster. California and Baja California 

vulnerata ( p. 130 ) 

Pronotum not strongly angulate at sides; color pattern not as 
above 33 

33. Vertex orange or red 34 

Vertex black or metallic 40 

34. Pronotum elongate, orange, with a wide black median stripe. 
Southern end of the Central Valley of Calif ornia . hoppingi (p. 171) 
Pronotum not as above 35 

35. Elytra and under surface metallic green or blue; pronotum trans- 
verse; head and pronotum hairy; length, 5 mm.-12 mm. South- 
western California aeneipennis (p. 174) 

Elytra and under surface black, not at all metallic; pronotum not 
transverse 36 

36. Elytra essentially glabrous; under surface dark pubescent; prono- 
tum without calluses on disk 37 

Elytra clothed with semierect, conspicuous silver pubescence (pro- 
ducing a dusty effect macroscopically ) ; average distance between 
setae near lateral margin of elytra about equal to length of a single 
seta; under surface pale pubescent; pronotum with a pair of large, 

strongly elevated calluses on disk. Southern Mexico 

proetus (p. 97 ) 

37. Antennal segments IV-X triangular (males, Fig. 91) or subtriangu- 
lar (females), produced on one side apically; under surface clothed 
with long shaggy pubescence; black with red head and pronotum. 

Southern Baja California Sur peninsularis (p. 141) 

Antennae, under surface, and distribution not as above 38 

38. Pubescence pale on head, pronotum, and under surface; pronotum 
campanuliform, one-tenth longer than wide; elytra piceous rather 
than a true black; tarsal pads hardly differentiated from rest of 
tarsal pubescence; male antennae without asymmetrically distorted 

segments ( Fig. 64 ) . Southern California and Baja California 

nitidicollis (p. 54 ) 

Pubescence dark; pronotum not as above; elytra a true black; tarsal 
pads distinct; male antennae with segments IV and V asymmetri- 
cally distorted (Figs. 95-96) 39 



40 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

39. Pronotum immaculate; antennal callosities and lower margin of 
frontal area of head black. Coahuila, Durango, Baja California 
Norte, extreme southern California, Arizona, and New Mexico. . . 

mutilata ( p. 152 ) 

Pronotum with a pair of large black spots on disk at middle (very 
rarely absent); entire front of head (not including epistoma) 
orange. Southern Sonora to Jalisco and Colima; thence southwest 
to Veracruz and Guerrero erythrothorax (p. 150) 

40. Head entirely black, without an orange frontal spot. Mexico .... 41 
Head black or metallic with an orange frontal spot 44 

41. Elytra metallic bluish purple or green. Veracruz, Guerrero, and 

Oaxaca bipiincticollis (p. 82) 

Elytra black 42 

42. Head and pronotum coarsely, densely punctate; elytra sparsely 
pubescent; first segment of female hind tarsi with some pale setae 

beneath at apex 43 

Head and pronotum finely, very sparsely punctate; elytra glabrous; 
first segment of female hind tarsi entirely black pubescent. Michoa- 
can and Guerrero michoacanae (p. 70) 

43. Antennal segments short, in part wider than long; male head be- 
hind eyes at most very shallowly concave. Southern Chihuahua to 

southwestern Mexico (state) and Puebla variabilis (p. 63) 

Antennal segments more elongate, none wider than long; discal 
pronotal spots very small, faint; male head behind eyes deeply 
concave. Nuevo Leon scituloides (p. 67) 

44. Pronotum orange with a pair of black spots on disk at middle, each 
spot on or covering a smooth, impunctate callus. Mexico (primarily 

southern), exclusive of Baja California 45 

Pronotum not as above. California and Baja California 47 

45. Elytral pubescence short, very sparse, inconspicuous, usually black; 

male pygidium strongly, narrowly produced ( Fig. 209 ) 

quadrimaculata (p. 93 ) 

Elytral pubescence moderately long, conspicuous; male pygidium 
not produced as above 46 

46. Pubescence on head pale golden; male head behind (and above) 

eyes deeply concave scitula (p. 90) 

Pubescence on head piceous; male head behind eyes not concave 
proteus (p. 97 ) 

47. Pronotum decidedly elongate, orange with a wide black median 

stripe hoppingi ( p. 171 ) 

Pronotum not as above 48 



SYSTEMATICS AND PHYLOGENY 41 

48. Metallic green or bluish green 49 

Black, without metallic luster 50 

49. Pronotum with a large green median spot; elytra with long pubes- 
cence confined to base. Fresno and Kern counties, California. . . . 

refulgens ( p. 175 ) 

Pronotum entirely orange; elytra clothed throughout with long, 
erect pubescence. Los Angeles County to San Diego County, Cali- 
fornia crotchi ( p. 177) 

50. Pronotum hexagonal (Fig. 190); male hind trochanters armed with 

a ventral spine at apex. Central Valley of California 

molesta ( p. 216 ) 

Pronotum subquadrate (Fig. 191); male hind trochanters not 

spined. Central California south to Baja California Sur 

childi (p. 218) 

51. Head entirely black or metallic, lacking even a pale frontal spot. .52 
Head with at least a small orange or red frontal spot, sometimes 
with orange or red on vertex also 65 

52. Elytra each with three distinct costae, these producing an undulate 

surface. Durango canelos (p. 79) 

Elytra without costae, not undulate 53 

53. Pronotum elongate, one-fifth longer than wide; male intermediate 

antennal segments not incrassate. Southwestern California 

lugens (p. 172) 

Pronotum not as above; not occurring in California 54 

54. Males 55 

Females 60 

55. Sixth abdominal sternum shallowly emarginate; lateral lobes of 

sternum not produced ( Figs. 225 and 227 ) 56 

Sixth abdominal sternum deeply emarginate; lateral lobes of ster- 
num produced ( Figs. 228, 229, and 231 ) 58 

56. Elytral pubescence short, sparse, not noticeable macroscopically. 

Southern Mexico 57 

Elytral pubescence long, dense, producing a dusty effect macro- 
scopically. Southwestern United States puberula (p. 68) 

57. Head and pronotum more or less coarsely, densely punctate; elytra 

at least sparsely pubescent, distinctly rugose variabilis (p. 63) 

Head and pronotum finely, sparsely punctate; elytra essentially 
glabrous, obsolescently rugose, nearly smooth. Tlaxcala and Puebla 
to Tabasco ebenina (p. 72 ) 



42 BIONOMICS, SYSTEM ATICS, AND PHYLOGENY OF LyttCl 

58. Lateral lobes of sixth abdominal sternum divergent (Fig. 231); 

pronotum relatively smooth augusti (p. 80) 

Lateral lobes of sixth abdominal sternum not appreciably diver- 
gent (Figs. 228-29) 59 

59. Elytra conspicuously clothed with moderately long, moderately 
dense pubescence; emargination of sixth abdominal sternum 
U-shaped, extending more than half length of sternum (Fig. 229) 

koltzei (p. 76 ) 

Elytra essentially glabrous; emargination of sixth abdominal ster- 
num broadly triangular, extending at most one-third length of 
sternum ( Fig. 228 ) plumb ea (p. 74 ) 

60. Elytral pubescence long, dense, producing a dusty effect macro- 
scopically; black, without metallic luster. Southwestern United 

States puberula (p. 68 ) 

Elytral pubescence not as above. Southern Mexico 61 

61. Sixth abdominal sternum truncate; black, without metallic luster 

62 

Sixth abdominal sternum shallowly, triangularly emarginate at apex 
(Figs. 275-76) 63 

62. Head and pronotum coarsely or moderately coarsely, densely punc- 
tate; elytra rugose, not shinier than head and pronotum, sparsely 
but clearly pubescent; first segment of hind tarsi with some pale 
setae beneath at apex; antennae as in Figure 121. Southern Chi- 
huahua to southwestern Mexico ( state ) and Puebla 

variabilis (p. 63 ) 

Head and pronotum finely, sparsely punctate, finely granular, dull; 
elytra nearly smooth, essentially glabrous, shinier than head and 
pronotum; first segment of hind tarsi entirely black pubescent; 
antennae as in Figure 122. Tlaxcala and southern Puebla to Tabasco 
ebenina (p. 72 ) 

63. Distal antennal segments as wide as or wider than long ( Fig. 126 ) ; 
antennae short, not reaching middle of pronotum; pronotum rela- 
tively smooth. Oaxaca, Veracruz, and Puebla augusti (p. 80) 

Distal antennal segments longer than wide; antennae longer, reach- 
ing basal third of pronotum; pronotum strongly alutaceous 64 

64. Elytra metallic dark blue or bluish green, moderately densely 
clothed with pale pubescence; pronotum not transversely oval ( Fig. 

171). Central Michoacan to eastern Puebla koltzei (p. 76) 

Elytra black with a tinge of dark blue, essentially glabrous, with a 
few black setae at base and apex; pronotum transversely oval (Fig. 



SYSTEMATICS AND PHYLOGENY 43 

170). Guanajuato and Michoacan to Distrito Federal 

plumbea (p. 74 ) 

65. Upper surface essentially glabrous; vertex or at least tempora 
orange or red; male antennal segments IV-VI distorted; length, 

14 mm.-32 mm. Coahuila and Durango to Panama 

eucera ( p. 144 ) 

Without the above combination of characters 66 

66. Southern Mexico ( Nayarit and Guanajuato southward ) 67 

United States, Baja California, and the northern tier of Mexican 
states 69 

67. Upper surface essentially glabrous; all clothing setae of body black. 

Michoacan and Mexico ( state ) erebea ( p. 143 ) 

Upper surface conspicuously pubescent; at least elytral pubescence 
pale 68 

68. Pubescence pale golden on head; male head behind (and above) 

eyes deeply concave scitiria (p. 90) 

Pubescence piceous on head; male head behind eyes not concave 
proteus ( p. 97 ) 

69. Vertex or at least tempora orange 70 

Vertex black or metallic 71 

70. Elytra glabrous, black. Central and southern California 

tenebrosa ( p. 116 ) 

Elytra clothed with long, fine pubescence, usually metallic green 
or bluish green. Southern California, Baja California, Arizona, and 
southern Utah auriculata (p. 165) 

71. Mandibles nearly straight; tarsal pads (pale pubescence on under 
surface of tarsi) completely absent; entirely black except for pale 

frontal spot. California funerea (p. 126) 

Mandibles strongly curved; tarsal pads present on at least fore and 
middle legs 72 

72. Males 73 

Females 94 

73. First fore tarsal segment bent, somewhat twisted, with a deep, 
socket-like excavation anteroventrally at base; bright metallic 
green, blue, purple, or violaceous; intermediate antennal segments 

distinctly incrassate; pronotum hexagonal 74 

First fore tarsal segment not so modified; lacking combination of 
other characters listed above 76 

74. Antennal segments IV-VIII flattened and clothed with long, erect 



44 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

setae on one side viridana ( p. 213 ) 

Antennal segments IV- VIII not so modified 75 

75. Elytra generally violet, at least on margins, rarely entirely green; 
pubescence on under surface of thorax pale; lateral lobes of fifth 

abdominal sternum angulate ( Fig. 219 ) nuttalli ( p. 205 ) 

Elytra lacking violet color; pubescence on under surface of thorax 
piceous; lateral lobes of fifth abdominal sternum rounded (Fig. 
220 ) cyanipennis ( p. 209 ) 

76. Fore tibiae each with a single spur 77 

Fore tibiae each with two spurs 78 

77. Antennae clavate (Fig. 110); hind trochanters angulate on ventral 

margin, not spined. Oregon and California moerens ( p. 196 ) 

Antennae with intermediate segments incrassate, in part as wide as 
distal segments; hind trochanters each armed with a distinct spine 
on ventral margin. Wyoming and Colorado. . .nigrocyanea (p. 202) 

78. Intermediate antennal segments incrassate, in part as wide as or 

wider than distal segments; black without metallic luster 79 

Intermediate antennal segments not incrassate, narrower than distal 
segments 85 

79. Hind trochanters each armed with a distinct spine on ventral margin 
(Fig. 162); sixth abdominal sternum extremely elongate, deeply 

cleft medianly ( Fig. 266 ) . Coastal southern California 

insperata ( p. 201 ) 

Hind trochanters not spined; sixth abdominal sternum not elongate, 
triangularly emarginate 80 

80. Middle tibiae each with a deep, impunctate, glabrous impression on 
anterior side apically; pygidium distinctly produced. Southern Cali- 
fornia 81 

Middle tibiae unmodified; pygidium not produced 82 

81. Pronotum as in Figure 181; pygidium blunt at apex (Fig. 213); 
lateral lobes of sixth abdominal sternum not slender ( Fig. 244 ) . . . 

melaena ( p. 122 ) 

Pronotum as in Figure 182; pygidium very acute at apex (Fig. 214); 
lateral lobes of sixth abdominal sternum produced as slender proc- 
esses ( Fig. 245 ) morosa ( p. 124 ) 

82. Antennal segments IV-VII with erect pale setae on ventral side. . . 

tenebrosa ( p. 116 ) 

Antennal segments IV-VII not so modified 83 

83. Pronotum subquadrate (Fig. 191); mesal surface of fore tibiae 
densely lined with golden pubescence; lateral lobes of sixth ab- 



SYSTEMATICS AND PHYLOGENY 45 

dominal sternum curving ventrad at apex, not hemispherical (Fig. 

270) childi (p. 218) 

Pronotum subhexagonal, with obtuse lateral angles (Figs. 179-80); 
fore tibiae not as above; lateral lobes of sixth abdominal sternum 
produced, hemispherical ( Fig. 243 ) 84 

84. One or both spurs on each hind tibia flattened, sticklike (Fig. 

195); wings pale brown; genitalia as in Figure 318 

bloisdelli ( p. 119 ) 

Both spurs on each hind tibia obliquely truncate (Fig. 194); wings 
colorless lugubris ( p. 120 ) 

85. Hind trochanters each armed with a distinct spine near apex. Cen- 
tral Valley of California molesta (p. 216) 

Hind trochanters not spined 86 

86. Hind trochanters angulate on ventral margin 87 

Hind trochanters not angulate 88 

87. Head and pronotum sparsely punctate; lateral lobes of sixth ab- 
dominal sternum not divergent ( Fig. 264 ) ; posterior fore tibial spur 

often much reduced in size. Oregon and California 

moerens ( p. 196 ) 

Head and pronotum densely punctate; lateral lobes of sixth ab- 
dominal sternum strongly divergent (Fig. 265). Northern Arizona 
navojo ( p. 200 ) 

88. Black, with dark metallic blue or green elytra; conspicuously clothed 
throughout with rather long, more or less erect pubescence; sixth 

abdominal sternum as in Figure 255 89 

Not as above 90 

89. Pubescence colorless apically or entirely colorless, at least on under 
surface of body; genitalia as in Figure 332. Southern California, 
Baja California, Arizona, and southern Utah. . .auriculata (p. 165) 
Pubescence piceous; genitalia as in Figure 333. Sierra Nevada and 
northern coastal region of California nigripilis (p. 169) 

90. Pronotum elongate, one-fifth longer than wide; black. Southwestern 

California lugens ( p. 172 ) 

Pronotum not elongate 91 

91. Elytra violet or green with a violet luster (not brassy); rest of body 
green or brassy; pubescence colorless. Sacramento and Calaveras 

counties, California, south to Tulare County, California 

rathvoni ( p. 178) 

Not as above 92 

92. Head and pronotum distinctly microreticulate, more or less satiny 



46 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LlfttCl 

in texture; sixth abdominal sternum relatively flat, shallowly, evenly 
emarginate (Fig. 262); antennal segments elongate, more cylin- 
drical (Fig. 106); pubescence colorless (Kern and Tulare counties, 

California ) or piceous ( central western California ) 

chloris ( p. 179 ) 

Head and pronotum smooth or relatively so; sixth abdominal ster- 
num medianly impressed, moderately deeply, triangularly emargi- 
nate; antennal segments more moniliform; pubescence piceous. .93 

93. Hairy, conspicuously clothed throughout with long, straight, erect 
pubescence; green. Central western California . . . comans ( p. 192 ) 
Pubescence shorter, less erect, especially on upper surface; elytral 
pubescence very short, subrecumbent, and inconspicuous, or if 
longer, erect setae present, these confined to immediate base; 
green, blue, or black. Idaho, Washington, Oregon, California, and 
southern Arizona stygica (p. 185) 

94. Black with dark metallic blue or green elytra; conspicuously clothed 
throughout with long, more or less erect pubescence; sixth ab- 
dominal sternum as in Figure 282 95 

Not as above; either black throughout or metallic throughout. .96 

95. Pubescence colorless apically or entirely colorless, at least on under 
surface of body. Southern California, Baja California, Arizona, and 

southern Utah auriculata (p. 165) 

Pubescence piceous. Sierra Nevada and nothern coastal region of 
California nigripilis (p. 169) 

96. Sixth abdominal sternum with a narrow, U-shaped emargination 
(Fig. 287); pronotum subquadrate; black. California and Baja 

California childi (p. 218) 

Sixth abdominal sternum without such an emargination 97 

97. Pronotum elongate, one-fifth longer than wide; black. Southwestern 
California (San Bernardino and Los Angeles counties to San Diego 

County ) lugens ( p. 172 ) 

Pronotum not elongate 98 

98. Head and pronotum very densely punctate, very densely clothed 
with long, erect, silky pubescence; black with or without a dark 
blue luster; elytra dark blue or indigo. Wyoming and Colorado. . . 

nigrocyanea ( p. 202 ) 

Not as above; if within range of nigrocyanea, then bright metallic 
in color 99 

99. Elytra coarsely, strongly rugose (subreticulate), dull; black with a 
metallic bluish luster; head and pronotum deeply, densely punctate. 
Northern Arizona navajo ( p. 200 ) 



SYSTEMATICS AND PHYLOGENY 47 

Elytra finely rugose, no more strongly so than usual for genus; if 
black with a metallic luster (some moerens), then head and pro- 
notum not densely punctate and range not including northern Ari- 
zona 100 

100. Outer hind tibial spur cylindrical, four times as wide as inner 
spur; black. Southwestern California and the Central Valley . . . 101 
Outer hind tibial spur twice or less than twice as wide as inner spur 
102 

101. Pale frontal spot at least half as wide as frontal area between eyes; 
sixth abdominal sternum emarginate (Fig. 286). Central Valley of 

California molesta (p. 216) 

Pale frontal spot at most one-fourth as wide as frontal area between 
eyes; sixth abdominal sternum entire. Ventura County to San Diego 
County, California insperata (p. 201 ) 

102. Bright metallic green, blue, purple, or violaceous 103 

Nonmetallic black, or black with metallic bluish luster (most moe- 
rens and some stygica ) 109 

103. Elytra violet or green with a violet luster; hind trochanters 
smoothly rounded on ventral margin; pubescence colorless through- 
out. Sacramento and Calaveras counties, California, to Tulare 

County, California rathvoni (p. 178) 

If elytra violet hind trochanters angulate on ventral margin, pu- 
bescence piceous on upper surface of body, and pronotum strongly 
hexagonal 104 

104. Elytra violet, at least on margins; hind trochanters angulate; pu- 
bescence pale on under surface of body. Alberta and Minnesota to 
Arizona and New Mexico, with a disjunct population in the Sierra 

Nevada of California nuttalli (p. 205) 

Elytra lacking violet color 105 

105. Pronotum distinctly hexagonal, with well-marked lateral angles (as 
in Fig. 189); apex of fore tibiae flared on posterior side; sixth 
abdominal sternum moderately deeply emarginate; hind trochan- 
ters often angulate on ventral margin 106 

Pronotum subhexagonal to nearly quadrate, without well-marked 
lateral angles; fore tibiae not so modified; sixth abdominal sternum 
not or only shallowly emarginate; hind trochanters never angulate 
on ventral margin 107 

106. Individual setae on under surface entirely piceous; hind trochanters 
often angulate. Southern British Columbia to northern California, 
southern Utah, and southern Wyoming cyanipennis (p. 209) 



48 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Individual setae on under surface colorless apically, piceous 
basally; hind trochanters never angulate. Peace River region of 
British Columbia and southern Manitoba to eastern Wyoming, 
Colorado, southeastern Utah, northern New Mexico, and nothern 
Arizona. Strictly allopatric with cyanipennis viridana (p. 213) 

107. Head and pronotum distinctly microreticulate, more or less satiny 
in texture; brilliant brassy green; pubescence colorless (Kern and 
Tulare counties, California) or piceous (central western Cali- 
fornia ) chloris ( p. 179 ) 

Head and pronotum smooth or relatively so; pubescence piceous 
108 

108. Hairy, conspicuously clothed throughout with long, straight, erect 
pubescence; green. Central western California. . .comans (p. 192) 
Pubescence shorter, less erect, especially on upper surface; elytral 
pubscence very short, subrecumbent, and inconspicuous, or if 
longer, erect setae present, these confined to immediate base; 
green or blue. Idaho, Washington, Oregon, California, and southern 
Arizona stygica ( p. 185 ) 

109. First segment of middle tarsi densely pale pubescent (padded) be- 
neath; first segment of hind tarsi also pubescent beneath, but 
pubescence pale only in apical part of segment; often with a metallic 
luster 110 

First segment of both middle and hind tarsi glabrous beneath be- 
tween lateral fringing bristles (completely lacking pads); never 
with a metallic luster Ill 

110. Tarsal claws heavy, strongly curved (Fig. 203); pubescence on 

under surface of body often pale. Oregon and California 

moerens ( p. 196 ) 

Tarsal claws slender, less strongly curved (Fig. 202); pubescence 
on under surface of body always piceous. Washington, Oregon, and 
northern California stygica ( p. 185 ) 

111. Head and pronotum granular, coarsely punctate, alutaceous; pro- 
notum subhexagonal, with obtuse lateral angles (Figs. 178-80) 

112 

Head and pronotum smooth, finely, sparsely punctate; pronotum 
without lateral angles ( Fig. 181 ) 114 

112. Hind tibial spurs distinctly dissimilar, the outer spur twice as wide 
as inner spur and obliquely truncate, and the inner spur flattened; 
pronotum widest near anterior margin (Fig. 178). Yolo and Marin 

counties, California, to San Diego County, California 

tenebrosa (p. 116) 



SYSTEMATICS AND PHYLOGENY 49 

Hind tibial spurs similar in size and often in shape; pronotum more 
nearly hexagonal, widest near middle (Figs. 179-80). Idaho, Ore- 
gon, and the Sierra Nevada of California 113 

113. Sixth abdominal sternum with hind margin turned gradually 
dorsad; one or both spurs of each hind tibia flattened, sticklike 

(Fig. 195) bloisclelli (p. 119) 

Sixth abdominal sternum with hind margin turned abruptly dorsad, 
at a right angle; both spurs of each hind tibia obliquely truncate 
( Fig. 194 ) lugubris ( p. 120) 

114. Pronotum somewhat quadrate (Fig. 181), as wide as long; fore 
tarsi nearly as long as fore tibiae. Southern California and the 

Yuma, Arizona, region to the tip of Baja California 

melaena ( p. 122 ) 

Pronotum subcircular, one-tenth or more wider than long; fore 
tarsi four-fifths as long as fore tibiae. In and near costal ranges in 
extreme southern California morosa (p. 124 ) 

Subgenus PARALYTTA, new subgenus 

Small to moderately large beetles. Colors usually nonmetallic; a dis- 
tinct metallic luster in some Mexican species, but color never brilliantly 
metallic throughout. Head when black with or without a pale frontal 
spot. Male antennae a little longer than in female, reaching at most three 
segments beyond base of pronotum; male intermediate antennal seg- 
ments incrassate, with at least IV and V fully as wide as or wider than 
distal segments, or if intermediate segments narrower, then IV wider 
than V (and IV and V usually wider than VI and VII); incrassate seg- 
ments somewhat compressed and usually flattened to some extent on 
inner side but never asymmetrically produced or excavate. Female an- 
tennae clavate. Elytral surface rugose or reticulate. First segment of male 
middle tarsi weakly distorted in a few species and male middle tibiae 
subapically impressed in a few others; male legs not otherwise modified. 
Each tibia with two spurs. Hind tibial spurs generally strongly dis- 
similar (exceptions in blaisdelli and lugubris), with outer spur much 
wider than inner spur and obliquely truncate and inner spur excavate 
and only slightly thickened. Male genitalia with gonostyli lacking mesal 
hooks, glabrous, tapered; aedeagus usually with two well-developed 
ventral hooks, rarely with only one. 

Type species. Lytta magister Horn. 

Remarks. This is the largest of the subgenera of Lytta, with 36 species, 
divided into five groups. It is restricted in distribution to the North 
American Continent. Its range extends from British Columbia, Canada, 
through the western half of the United States, to southern Mexico. Most 



50 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lljttd 



of the species found in the United States occur also in Mexico, while 
16 of the species are strictly Mexican in distribution. On the basis of both 
adult and larval morphology, the subgenus most closely resembles and 
is probably most closely related to the subgenus Adicolytta. 

The majority of host plants recorded for species of Paralytta belong 
to the families Compositae, Convolvulaceae, and Papaveraceae. Other 
families represented by dependable records are Boraginaceae, Campanu- 
laceae, Leguminosae, Liliaceae, Scrophulareaceae, Labiatae, and Pri- 
mulaceae. According to my field observations on eight species of the 
Variabilis Group, the principal food of the adults of this group, and 
perhaps of the other groups as well, is pollen. Petals are sometimes eaten, 
but I have never observed feeding on the foliage of the host plants. 
Larval hosts have been recorded for three species (Table I). 

The relationships of the species of Paralytta suggested by my study 
are shown in Figure 10 and are discussed in the following paragraphs. 
It will be evident that I have based my phylogeny on static relationships 
to a large extent in this subgenus. 

The Variabilis and Reticulata groups are sufficiently alike that they 
might reasonably be combined as a single group. In the males of all 
species of both groups the surface of the head behind the eyes is 
densely micropunctate and finely punctate, and the setae in this region 
are very short. In most members of the Variabilis Group this area is 
also either flattened or impressed. This modification of the head is un- 




Fig. 10. Phylogeny of the subgenus Paralytta. 



SYSTEMATICS AND PHYLOGENY 



51 



doubtedly a specialized character, and for this reason the two groups 
presumably belong to a distinct phyletic line. The Reticulata Group is 
further specialized in having reticulate, rather than rugose, elytra. It 
cannot, to my way of thinking, be derived from any of the three sub- 
groups of the Variabilis Group; it is therefore necessary to postulate that 
the two groups diverged from an early ancestral type. In the Variabilis 
Group line the pronotal disk has developed a pair of smooth calluses, 
which is present in all species except ebenina. The Variabilis Group pro- 
liferated primarily in central and southern Mexico; only two of its 
species (biquttata and puberula) are found as far north as the northern 
border of Mexico. On the other hand, the Reticulata Group has its dis- 
tribution centered in the southwestern United States and adjacent north- 
ern Mexico, with an extension onto the Great Plains (reticulata) and 
another into central Mexico ( corallifera ) . 

Within the Variabilis Group the Variabilis and Koltzei subgroups may 
be derived as branches of a line which lost the pale frontal spot of the 
head while retaining or largely retaining the essentially black coloration 
which I believe is primitive for the subgenus. In the Quadrimaculata 
Subgroup the spot has been retained, and there has been a well- 
developed trend for the development of pale colors on the body and legs. 
In addition, all the species of the subgroup except sonorae have a char- 
acteristic distortion of the first segment of the male middle tarsi. A some- 
what similar modification is found in two species of the Koltzei Subgroup 
(augusti and bipuncticollis), apparently as the result of convergent 
evolution. Within the Reticulata Group two main lines may be distin- 
guished on the basis of the presence or absence of the frontal spot 
(absent in the Reticulata Subgroup) and on the basis of a difference in 
the general form of the male genitalia. 

The Magister Group represents a distinct line which apparently under- 
went most of its evolutionary development in the far-western United 
States, well to the north and west of the presumed evolutionary centers 
of the Variabilis and Reticulata groups. The Tenebrosa Subgroup is an 
essentially montane offshoot of the main line of the group which appar- 
ently had an early origin in view of its morphological distinctness and 
the lack of modification of the male pygidium. The other two subgroups 
agree in having the pygidium produced and in having the male middle 
tibiae at least weakly distorted. There is also a tendency for reduction 
of the tarsal pads in these two subgroups, the pads having been lost 
independently in magister and funerea. 

Taken as a whole, the Magister Group is much more sharply defined 
morphologically in the larval stage than in the adult. According to Mac- 
Swain (1956, p. 78), who has described the larvae of five of the nine in- 
cluded species, the Magister Group in the larval stage is distinguished 



UNIVERSITY OF ILLINOIS. 
LIBRARY 



52 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

from all other Lyttini by the following characters: "maxillary palpus with 
three or more long setae on anterior dorsal margin of second segment, 
setae longer than third segment; first two segments of maxillary palpi un- 
equal, well sclerotized, broad." These characters are assuredly specializa- 
tions, and since the group in the adult stage is more primitive than the 
Variabilis and Reticulata groups in several characters, including the 
absence of modification of the male head, it is reasonable to conclude 
that the group diverged from the paralyttine stock before the evolution 
of the lines forming the Variabilis and Reticulata groups. 

The Fulvipennis Group closely approaches the Variabilis Group but 
does not have the head of the male modified. It might conceivably repre- 
sent an offshoot of the M agister Group line, but there is nothing in par- 
ticular to suggest this in the adult morphology. Unfortunately, larval 
morphology cannot at present be brought to bear on this problem since 
the larvae of fulvipennis and lecontei are unknown. Pending additional 
information, I regard the Fulvipennis Group as a primitive line which 
evolved independently of the other groups of the subgenus Paralytta. 

The Nitidicollis Group contains three Californian species of uncertain 
relationships. The species nitidicollis is a form which on the basis of 
both adult and larval morphology cannot be satisfactorily placed within 
any of the other groups of the subgenus but which seems to lack any 
obvious specialization, except possibly with regard to adult coloration. 
In the adult stage morrisoni is quite similar to nitidicollis with the im- 
portant exception that it has the two blades of the tarsal claws fused 
basally, as in the members of the tribe Eupomphini. The tarsal claws of 
moesta are modified in the same manner, but this species is otherwise 
distinct enough from both nitidicollis and morrisoni that it is a little diffi- 
cult to believe that the modification of its claws and those of morrisoni 
stems from a common ancestor. On the contrary, were it not for the 
modified claws and the lack of modification of the male middle tibiae, 
moesta would fit nicely into the Magister Group next to funerea. It is 
hoped that larval studies will help to resolve the difficulties here out- 
lined. 

Key to Groups 

1. Tarsal claws with ventral blade fused basally with dorsal blade and 

much shorter than it Nitidicollis Group ( part ) ( p. 53 ) 

Tarsal claws cleft to base, with blades subequal in length 2 

2. Male head behind eyes more densely micropunctate, more finely 
punctate, and usually duller than at center of vertex and clothed 
with distinctly shorter setae; this region also frequently flattened 
or impressed; if elytra reticulate, male pygidium not produced; if 



SYSTEMATICS AND PHYLOGENY 53 

elytra rugose, pronotum usually with a pair of smooth calluses (ex- 
ception in ebenina ) 3 

Male head not modified as above (setae behind eyes shortened in 
nitidicollis, but surface sculpture normal ) 4 

3. Elytra rugose or rugose punctate; pronotum usually with a pair of 

calluses Variabilis Group (p. 61 ) 

Elytra strongly reticulate; pronotum without calluses 

Reticulata Group ( p. 101 ) 

4. Pronotum hexagonal or male pygidium triangularly produced, or 

both; elytra rugose or reticulate, black Magister Group (p. 113) 

Pronotum campanuliform or transversely oval, without lateral angles; 
pygidium not produced; elytra rugose, usually orange 5 

5. Pronotum broad, transversely oval Fulvipennis Group (p. 57) 

Pronotum narrow, campanuliform. .Nitidicollis Group (part) (p. 53) 

NITIDICOLLIS GROUP 

Colors nonmetallic. Head when black with a pale frontal spot. Head 
quadrate; tempora broadly rounded, not prominent; surface smooth, 
sparsely micropunctate, finely punctate, subglabrous. Male head not 
modified behind eyes, except that setae in this area are shortened in 
nitidicollis. Male antennae reaching about base of pronotum; male inter- 
mediate segments ( IV-VII ) weakly incrassate. Pronotum campanuli- 
form, longer than wide; surface similar to that of vertex. Elytra rugose. 
Legs unmodified. Tarsal pads poorly differentiated from rest of tarsal 
pubescence. Tarsal claws either cleft to base and with blades subequal 
in length (nitidicollis) or with ventral blade fused basally with dorsal 
blade and much shorter than it. Male sixth abdominal sternum shallowly 
emarginate. Male pygidium produced (moesta) or not. Aedeagus with 
two ventral hooks. 

Remarks. As mentioned above, this group is probably an unnatural 
one. Included in it are the species nitidicollis, morrisoni, and moesta. The 
last two of these species differ from all other Lyttini in having the two 
blades of the tarsal claws fused, as in the members of the tribe 
Eupomphini. All three species have a similar general facies and exhibit 
certain peculiarities of the tarsal pubescence. None can be placed in 
any of the other groups of Paralytta without making major changes in the 
definitions of these groups. 

The species morrisoni and moesta have essentially the same range, in 
central California (Fig. 5). Nitidicollis is found in southern California 
and on the peninsula of Baja California. 



54 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Key to Species 

1. Tarsal claws cleft to base, with blades subequal in length; head, 

pronotum, and ( usually ) elytra orange nitidicollis ( p. 54 ) 

Tarsal claws with ventral blade fused basally with dorsal blade and 
much shorter than it; at least head and elytra black 2 

2. Elytra moderately densely clothed with semierect setae; tarsal claws 
distinctly curved; emargination of male sixth abdominal sternum 
sparsely pubescent; male pygidium not produced; pronotum often 
orange morrisoni ( p. 55 ) 

Elytra essentially glabrous; tarsal claws nearly straight; emargina- 
tion of male sixth abdominal sternum densely lined with short setae; 
male pygidium produced; pronotum always black. . . .moesta (p. 56) 

Lytta (Paralytta) nitidicollis (LeConte) 

Cantharis nitidicollis LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 160. Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. 112; 1883, Trans. 
Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Occas. Papers California Acad. 
Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc., vol. 27, p. 299. 

Lytta nitidicollis, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 332. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 84, pi. 15. 

Black. Head and pronotum rich brownish or reddish orange. Elytra 
and scutellum either a similar (but often duller) orange or (rarely) 
piceous. Wings light brown. Pubescence yellow on head, pronotum, and 
elytra (unless these last are piceous), silver on under surface of body, 
piceous or nearly so on legs. Length: 12 mm. -17 mm. 

Head sparsely punctate. Pronotum one-tenth longer than wide; sur- 
face nearly always weakly alutaceous but otherwise as on vertex. Elytra 
weakly rugose, very finely, obscurely punctate, essentially glabrous. 
Ventral surface of first segment of middle and hind tarsi densely pubes- 
cent, but pubescence hardly differentiated (except occasionally at apex 
of segment) from rest of pubescence of segment; other tarsal segments 
with better developed, undivided, pale pads. Tarsal claws slender, weakly 
curved, cleft to base, with blades subequal in length. 

Male. Head with setae behind eyes somewhat shortened, but integu- 
ment in this region not modified. Antennae as in Figure 64; segments a 
little more elongate than in morrisoni. Fifth abdominal sternum moder- 
ately deeply emarginate. Emargination of sixth sternum rounded, lined 
with short setae; lateral lobes of sternum subacute. Pygidium not pro- 
duced. Genitalia as in Figure 290. 

Female. Antennae as in Figure 117. Sixth abdominal sternum feebly 
emarginate medianly. 



SYSTEMATICS AND PHYLOGENY 



55 



Type locality. San Diego, California. 

Geographic distribution. Santa Clara County, California, south an 
unknown distance on the peninsula of Baja California. (See Fig. 11.) 

Seasonal distribution. May 29 — July 20. 

Records. UNITED STATES: California: State label only, 28; Dulzura, 
July, 1; Gavilan Peak, May, 4; Gilman Hot Springs, Riverside County, 
June, July, 16; La Jolla, July, 1; Pasadena, June, 1; Riverside (Mac- 
Swain, 1956); San Diego, 6 (4 in May); San Diego County, 1; Santa 
Clara County, 2. 

MEXICO: Baja California [Norte?]: (Fall, 1901, p. 299). 

Remarks. The elytra are entirely piceous in the specimen from La 
Jolla and in one of the two specimens from Santa Clara County, Cali- 
fornia. In the other specimen from Santa Clara County they are orange 
irregularly suffused with piceous. 

The larva of nitidicollis has been described by MacSwain (1956). 
The type is in the Museum of Comparative Zoology. 

Bionomics. The specimens from Gavilan Peak, California, are recorded 
from Salvia midtiflora (Labiatae). 

Lytta (Paralytta) morrisoni (Horn) 

Calospasta morrisoni Horn, 1891, Proc. Amer. Philos. Soc, vol. 29, p. 102. 
Wellman, 1909, Ent. News, vol. 20, p. 24. 

Lytta morrisoni, Selander, 1954, Coleopterists' Bull, vol. 8, p. 11. 




Fig. 11 (left). Distribution of Lytta nitidicollis. 
Fig. 12 (center). Distribution of Lytta morrisoni. 
Fig. 13 (right). Distribution of Lytta mocsta. 



56 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Black. Pronotum orange or black. Frontal spot elongate, triangular, 
one-third or less as wide as frontal area between eyes. Wings dull brown. 
Length: 12 mm. -17 mm. 

Head one-tenth longer than wide; surface moderately densely punc- 
tate. Eyes wider than in moesta. Pronotum slightly longer than wide; 
surface a little less densely punctate and pubescent than on vertex. Elytra 
impunctate, moderately densely clothed with short, semierect setae. 
Wings relatively broad. Ventral surface of first segment of tarsi densely 
clothed with pubescence which is not differentiated from that of rest of 
tarsi; rest of segments with pale pads, but the setae of these only slightly 
recumbent and not greatly different in texture from rest of tarsal pubes- 
cence. Tarsal claws distinctly curved; ventral blade fused basally with 
dorsal blade and about half as long as it. 

Male. Antennae as in Figure 65. Fifth abdominal sternum shallowly 
but obviously emarginate. Emargination of sixth sternum as in Figure 
222, sparsely pubescent; lateral lobes of sternum rounded. Pygidium not 
produced. Genitalia as in Figure 291; gonostyli cylindrical, hardly 
tapered, bluntly rounded at apex; gonocoxal piece truncate. 

Female. Antennae as in Figure 118. Sixth abdominal sternum entire. 

Type locality. Southern California. 

Geographic distribution. Central California. (See Fig. 12.) 

Seasonal distribution. April 15 — May 9. 

Records. UNITED STATES: California: Coalinga, May, 3; Edison, 
April, 10; Kaweah, 4; Kern County, 2; Southern California, 11; Tipton, 
April, 1; south of White River, April, 3. 

Remarks. The pronotum is entirely black in 2 of the 11 specimens 
labeled Southern California and in 2 of the 3 specimens from Coalinga. 
In the third specimen from Coalinga it is largely black but has some 
orange coloring. In all specimens from Edison, Kaweah, Tipton, and 
south of White River it is orange. 

The type is in the collection of the Academy of Natural Sciences of 
Philadelphia. 

Lytta (Paralytta) moesta (Horn) 

Calospasta moesta Horn, 1878, Trans. Amer. Ent. Soc, vol. 7, p. 59; 1891, 
Proc. Amer. Philos. Soc, vol. 29, p. 101. Wellman, 1909, Ent. News, vol. 20, 
p. 24. 

Lytta moesta, Selander, 1954, Coleopterists' Bull., vol. 8, p. 11. 

Black. Frontal spot elongate oval, one-third or less as wide as frontal 
area between eyes. Wings very pale yellowish brown, darker at apex. 
Length: 16 mm. -20 mm. 

Head slightly longer than wide; surface moderately densely punctate. 



SYSTEMATICS AND PHYLOGENY 57 

Pronotum elongate; surface a little more coarsely and densely punctate 
than on vertex, sometimes sparsely alutaceous. Elytra impunctate, essen- 
tially glabrous. Wings narrower than in morrisoni. Ventral surface of 
tarsi densely clothed with suberect black pubescence of about same 
color, length, and texture as rest of tarsal pubescence; hind tarsi some- 
times ventrally glabrous along midline. Tarsal claws nearly straight; 
ventral blade fused basally with dorsal one and usually less than half as 
long as it. 

Male. Fifth abdominal sternum entire or only feebly emarginate. 
Emargination of sixth sternum as in Figure 233, densely lined with 
short setae. Pygidium (Fig. 204) produced, blunt at apex. Genitalia as 
in Figure 292. 

Female. Pygidium not or only weakly produced. Sixth abdominal 
sternum entire. 

Type locality. California. 

Geographic distribution. Central California. (See Fig. 13.) 

Seasonal distribution. April 2 — September 15. 

Records. UNITED STATES: California: State label only, 2; Edison, 
April, 1; Friant, April, 1; Kaweah, 4; Kismet, September, 5; Manteca, 
August, 2; Modesta, September, 1; Potwisha, Sequoia National Park, 
2000 ft.-5000 ft., May, 1; Ripon, 3; Santa Cruz, April, 4; Tulare County, 
3; Westley, July, 1. 

Remarks. The type is in the collection of the Academy of Natural 
Sciences of Philadelphia. 

FULVIPENNIS GROUP 

Elytra immaculate yellow or orange. Pubescence pale on elytra, black 
elsewhere. Head subquadrate; tempora broadly rounded, not prominent; 
surface smooth, finely punctate. Male head not modified behind eyes. 
Male antennae reaching about middle of pronotum; male intermediate 
segments (IV- VII) weakly incrassate, subequal, as wide as or very 
slightly wider than following segments. Eyes broad, bulged. Pronotum 
transversely oval, without distinct calluses. Scutellum densely punctate 
and pubescent. Elytra rugose, dull, veiy finely, densely punctate, sub- 
glabrous. Legs unmodified. Outer hind tibial spur unusually thick, 
cylindrical, expanded distally, three times as wide as inner spur. Tarsal 
pads well developed, undivided; pale pubescence ( pad ) of first segment 
of middle and hind tarsi of female and hind tarsi of male limited to 
apex. Tarsal claws heavy, cleft to base, with blades subequal in length. 
Male fifth abdominal sternum moderately deeply emarginate. Emargina- 
tion of male sixth sternum shallow; a fine groove extending anteriad from 
apex of emargination; lateral lobes of sternum not produced. Female 



58 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

sixth sternum truncate, lacking groove. Pygidium not produced. Aede- 
agus with two ventral hooks. 

Remarks. This group contains two species: fulvipennis and lecontei. 
These species occur together in eastern Texas, including the Rio Grande 
Valley, with fulvipennis ranging west to eastern Arizona and north, on 
the Great Plains, to Nebraska (Fig. 5). 

Key to Species 

1. Head and pronotum black, densely punctate, and densely pubescent 

fulvipennis (p. 58 ) 

Head and pronotum orange or yellow (pronotum with two black 
spots), sparsely punctate, and sparsely pubescent. . .lecontei (p. 60) 

Lytta (Paralytta) fulvipennis LeConte 

Lytta fulvipennis LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 331; 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 447. Dillon, 1952, 
Amer. Midland Nat., vol. 48, p. 351, pi. 2, fig. 1. 

Cantharis cardinalis, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 112; 
1883, Trans. Amer. Ent. Soc., vol. 10, p. 311. Misidentifications. 

Cantharis fulvipennis, Horn, 1885, Trans. Amer. Ent. Soc, vol. 12, p. 111. 
Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 298. Wickham, 1902, Bull. Lab. 
Nat. Hist. State Univ. Iowa, vol. 5, p. 301. Pritchett, 1903, Biol. Bull., vol. 5, 
pp. 275, 279. 

Black. Elytra rich brownish yellow or orange. Head with a diamond- 
shaped orange frontal spot. Wings yellow, except apical fourth brown. 
Pubescence pale on elytra, piceous elsewhere. Length: 11 mm.-26 mm. 

Head with sides nearly parallel above eyes; surface moderately 
shiny, very finely, densely punctate; average distance between punctures 
about equal to diameter of a single puncture; pubescence moderately 
long, erect, moderately dense. Pronotum generally wider than long 
(rarely no wider than long), widest at middle; disk even, sometimes 
with a small, shallow impression on each side behind middle; surface 
as on head but more densely punctate and pubescent; midline and a few 
small, scattered areas on disk impunctate. 

Male. Antennae as in Figure 66; emargination of sixth abdominal 
sternum (Fig. 224) shallowly triangular. Genitalia as in Figure 293. 

Female. Antennae as in Figure 119. 

Type locality. Texas. 

Geographic distribution. Coastal Plain of Texas northwest to extreme 
southeastern Arizona (Chiricahua Mountains) and north on the Great 
Plains to southwestern Nebraska. (See Fig. 14.) The range undoubtedly 
extends some distance into Mexico, although there are no Mexican 
records at present. 



SYSTEMATICS AND PHYLOGENY 



59 



In addition to the material listed below, I have seen a series of 
five specimens in the United States National Museum labeled "near 
Hanksville, base of Henry Mountains, Utah, June 1936, E. Bowles." 
This locality is so distant from what may be considered the established 
range of the species that it seems likely that the specimens in question 
were accidentally mislabeled. 

Seasonal distribution. The earliest collection date for specimens I 
have examined is March 2; the latest, July 14. According to Dillon 
(1952), the species occurs in Texas as late as September. 

Records. UNITED STATES: Arizona: Cove Creek Canyon, Chirica- 
hua Mountains, July, 11. Colorado: (Horn, 1873). Kansas: Ashland, 
June, 4; Barker County, 1468 ft., 1; Belvidere, June, 2; Cheyenne County, 
3300 ft., 1; Clark County, 1962 ft., June, 8; Comanche County, 2089 ft, 
1; Kiowa County, 7 (5 in July); Rawlins County, July, 1; Sun City, June, 
3; Wellington, 1. Nebraska: Haigler, July, 1; Stratton, July, 4. New 
Mexico: Artesia, 1. Oklahoma: Lawton, 1; Rosston, 4; Quartz Mountain 
State Park, June, 43. Texas: State label only, 16; Abilene, June, 2; Austin, 
May, June, 5; Baby Head, May, 1; Baird, June, 1; Bastrop, 2; Bell 
County, June, 3; Big Spring, July, 12; Brownwood, June, 2; Burnet 
County, June, 1; Carrizo Springs, April, 1; Dallas, 4; Dilley (Dillon, 
1952); Eagle Pass, May, 1; Edinburg, April, 2; Fort Sam Houston, June, 
1; Gillispie County, June, 2; Hidalgo County, May, 1; Hillsboro, June, 




Fig. 14 (left). Distribution of Lijtta fuhipennis. 
Fig. 15 (right). Distribution of Lytta lecontei. 



60 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

8; Lamposas, June, 1; Laredo, April, 1; Luling, May, 5; McAllen, March, 
1; McLennan County, June, 2; Mason, May, 8; Mission, March, 4; New 
Braunfels, May, June, 30; Odessa, May, 1; Oklaunion, June, 13; Pease 
River, 2 mi. east of Vernon, 1000 ft.-1500 ft., July, 2; 5 mi. east of Rio 
Grande [City], April, 3; Round Mountain, 6; Sabinal, May, 1; San 
Antonio, May, June, 7; Sanderson, June, 2; Sequin, June, 3; Sweetwater, 
1; Tarrant County (Dillon, 1952); Taylor, April, May, 9; Temple, May, 
3; Victoria, May, 3; Waco, June, 4; Zephr, June, 3. 

Remarks. This species is easily recognized by its unique color pattern. 
It exhibits little in the way of individual variation and none of a geo- 
graphic nature. 

Three of the females examined have greatly enlarged heads. In two 
of them the head is strongly triangular, while in the third it retains 
the normal, more or less quadrate form. I have observed similar anomalies 
in a few specimens of several other species of Meloidae, among them 
Lytta erythrothorax. Two of the specimens of fulvipennis belong to the 
large series from Big Spring, Texas; the other specimen is recorded 
merely from Texas, without precise locality data. The type of fulvipennis 
is in the Museum of Comparative Zoology. 

Bionomics. According to label data, adults of fulvipennis have been 
collected on Helianthus and "thistle" (Compositae) in Texas and on 
Argemone platyceras ( Papaveraceae ) in Texas and Oklahoma. 

Lytta (Paralytta) lecontei Heyden 

Lytta dichroa LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 332. Not Fischer von Waldheim, 1823-24, Entomographie de la Russie et 
genres des insectes . . . , vol. 2, p. 230. 

Cantharis dichroa, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. Ill; 
1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. 
Soc, vol. 27, p. 298. 

Lytta lecontei Heyden, 1890, Wiener Ent. Zeitung, vol. 9, p. 99. Van Dyke, 
1929, Bull. Brooklyn Ent. Soc, vol. 24, p. 131. Dillon, 1952, Amer. Midland 
Nat, vol. 48, p. 352. 

Black. Vertex and upper frontal area, pronotum (except margins), 
and elytra rich brownish orange or yellow. Pronotum with a small 
black spot on each side of disk before middle. Wings yellow, with apex 
a little darker. Pubescence pale on elytra, black elsewhere. Length: 
13 mm. -20 mm. 

Head with sides weakly to moderately divergent above eyes; vertex 
shiny, finely, sparsely punctate, usually with a few micropunctures inter- 
mixed with others; frontal area (especially lower half) distinctly more 
densely, deeply punctate than vertex, with a shallow impression on each 
side or with a deep, continuous impression between eyes (in which case 
vertex appears tumid); pubescence short, erect. Pronotum one-tenth 



SYSTEMATICS AND PHYLOGENY 61 

wider than long, widest at middle; disk smooth, shiny, more irregularly 
punctate and pubescent than vertex; a shallow median impression at 
base. Elytral punctures not well defined. 

Male. Antennae as in Figure 67; anterior spur of fore and middle 
tibiae longer and stouter than posterior spur. Pale pubescence (pad) of 
first segment of middle tarsi limited to apex (as in female). Emargina- 
tion of sixth abdominal sternum shallowly oval. Genitalia as in Figure 
294. 

Female. Antennae as in Figure 120. 

Type locality. Boundary of the United States and Mexico. 

Geographic distribution. Eastern Texas and the lower end of the 
Rio Grande Valley (Texas and Tamaulipas). (See Fig. 15.) 

Seasonal distribution. March 20 — September 9. 

Records. UNITED STATES: Texas: State label only, 12; Alamo, 
April, 2; Bell County, June, 1; Bishop, April, 3; Brazos County, Septem- 
ber, 1; Brownsville, March, April, 4; Dallas, 2 (1 in May); Fort Worth, 
May, 4; Harlingen, March, 1; Kingsville, 1; McAllen, April, 1; McKinney, 
1; Taylor, May, 3. 

MEXICO: Tamaulipas: Rio Rico, August, 1. 

Remarks. The type of dichroa, and thus of lecontei, is in the Museum 
of Comparative Zoology. 

Rionomics. The three specimens from Bishop, Texas, are recorded as 
"feeding on primrose" ( Primulaceae ) . In addition, there are single rec- 
ords each from alfalfa and beans ( Leguminosae ) , Monarda citridosa 
(Labiatae), oats (Gramineae), and sunflower (Compositae), and two 
records from cotton (Malvaceae). At least some of these records un- 
doubtedly represent accidental associations. 

VARIABILIS GROUP 

Male head behind eyes more densely micropunctate, more finely 
punctate, and usually duller than at center of vertex, and clothed with 
much shorter (very short) setae; this region usually flattened or im- 
pressed. Male antennae with intermediate segments weakly to very 
strongly incrassate. Eyes broad, prominent. Pronotum oval, circular, or 
campanuliform; disk almost always with a smooth, impunctate callus on 
each side at or near middle (sometimes poorly developed; absent con- 
sistently only in ebenina). Elytra rugose or finely rugose reticulate; 
reticulations when present never clearly defined macroscopically. Outer 
hind tibial spur two to three (rarely four) times as wide as inner spur. 
First segment of male middle tarsi sometimes bent. Tarsal pads moder- 
ately to poorly developed, undivided. Tarsal claws cleft to base, with 



62 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ll/ttCl 

blades subequal in length. Pygidium produced or not. Aedeagus with 
two ventral hooks or (bipuncticollis and proteus) with only one. 

Remarks. This group contains 15 species, divided into three subgroups. 
It is primarily Mexican in distribution (Fig. 2). 

Key to Subgroups 

1. Head with at least a pale frontal spot; elytra usually partially or 
entirely pale; under surface and legs often pale, at least in part. . . . 

Quadrimaculata Subgroup (p. 84) 

Head entirely black, lacking a pale frontal spot; elytra, under sur- 
face, and legs entirely black or dark metallic 2 

2. Male sixth abdominal sternum shallowly emarginate; lateral lobes of 
sternum not produced; female sixth abdominal sternum truncate .... 

Variabilis Subgroup (p. 62) 

Male sixth abdominal sternum deeply emarginate; lateral lobes of 
sternum produced; female sixth abdominal sternum shallowly, trian- 
gularly emarginate at apex Koltzei Subgroup (p. 73) 

Variabilis Subgroup 

Black throughout (with or without a faint metallic luster) or with 
pronotum pale, bimaculate. Head lacking a pale frontal spot. First 
segment of male middle tarsi unmodified. Male fifth abdominal sternum 
feebly emarginate, almost entire. Male sixth sternum shallowly emargin- 
ate; lateral lobes of sternum not produced. Female sixth sternum trun- 
cate. 

Remarks. This subgroup contains the species variabilis, scituloides, 
puberula, michoacande, and ebenina. Of these, only puberula is known to 
occur north of the Mexican border. The first three species listed form 
a very homogeneous complex, while michoacanae and ebenina are more 
distinctly differentiated. The known host plants of the subgroup include 
species of Papaveraceae (ebenina), Convolvulaceae (variabilis, micho- 
acanae), and Campanulaceae (puberula). 

Key to Species 

1. Head and pronotum coarsely or moderately coarsely, densely punc- 
tate; elytra at least sparsely pubescent; first segment of female hind 

tarsi with some pale setae beneath at apex 2 

Head and pronotum finely, sparsely punctate; elytra essentially gla- 
brous; first segment of female hind tarsi entirely black pubescent. .4 

2. Elytral pubescence short, sparse, not noticeable macroscopically 
(average distance between setae greater than length of a single 
seta); pronotum black or orange. Mexico 3 



SYSTEMATICS AND PHYLOGENY 63 

Elytral pubescence relatively long, dense, conspicuous macro- 
scopically, producing a dusty effect (average distance between setae 
distinctly less than length of a single seta); pronotum black. South- 
western United States puberula (p. 68) 

3. Male head behind eyes deeply concave; antennal segments more 
elongate, none wider than long; when pronotum orange, discal spots 

very small, faint scituloides (p. 67) 

Male head behind eyes flattened or very shallowly concave; antennal 
segments short, in part wider than long (Fig. 68); pronotum black 
or orange with larger discal spots variabilis (p. 63) 

4. Pronotum orange; head and pronotum smooth, shiny; elytra distinctly 

rugose; pubescence pale on under surface michoacanae (p. 70) 

Pronotum black; head and pronotum finely granular, dull; elytra 

obsolescently rugose; pubescence black on under surface 

ebenina ( p. 72 ) 

Lytta (Paralytta) variabilis (Duges) 

Cantharis variabilis Duges, 1869, La Naturaleza, vol. 1, p. Ill, fig. 11; 
1889, An. Mus. Michoacano, vol. 2, p. 107. Champion, 1892, Biol. Centrali- 
Amerieana, Coleoptera, vol. 4, pt. 2, p. 447. 

Lytta variabilis, Selander, 1954, Jour. Kansas Ent. Soc, vol. 27, p. 90. 
Linslev, MacSwain, and Smith, 1956, Bull. Southern California Acad. Sci., 
vol. 55, pp. 93-95. 

Black (a little brassy in fresh material). Pronotum either orange with 
two black discal spots ( rarely small dots on each side at middle ) or com- 
pletely black. Wings light brown. Pubescence silver (except, often, on 
tibiae and tarsi), dark, or dark on upper surface and legs and silver 
on under surface, not conspicuous macroscopically. Length: 7.5 mm.- 
13.0 mm. 

Head with sides divergent above eyes; surface typically coarsely, 
deeply, densely punctate, with punctures crowded on frontal area and 
center of vertex, producing a rough surface; frontal area generally with a 
transverse, less densely punctate or impunctate region at center between 
eyes; pubescence moderately long, erect. Pronotum (Fig. 168) transverse, 
at least one-tenth wider than long; disk somewhat flattened, especially 
before middle, with a broad impression on midline at base; surface 
typically roughened by coarse, deep punctures like those on vertex; 
midline and pair of calluses impunctate; pubescence as on vertex but 
slightly shorter, tending to slant toward calluses on each side. Elytra 
rugose, lacking distinct punctures except for some fine ones near base; 
pubescence sparse, semierect; average distance between setae at middle 
of elytra at least equal to length of a single seta. Tarsal claws (Fig. 
199) heavy, very strongly curved. 



64 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



Fig. 16. Distribution of Lytta variabilis (circles) and Lytta scituloides (squares). 

Male. Head behind eyes flattened or shallowly concave. Antennae 
(Fig. 68) short, reaching middle of pronotum; segments III-V weakly 
incrassate; IV largest, as wide as VIII; V and VI similar in size; VI-VIII 
progressively wider; VIII-X subequal in width; segments short, VI-X 
as wide as or wider than long. Sixth abdominal sternum (Fig. 225) 
very shallowly emarginate, almost truncate. Pygidium with hind margin 
usually more oval in outline than in female but lacking a distinct apex. 
Genitalia as in Figure 295; aedeagus with hooks moderately variable 
in form. 

Female. Antennae (Fig. 121) definitely short of middle of pronotum; 
segments VI-X as in male. Pad of first segment of middle tarsi limited 
to apical half. 

Type locality. Guanajuato, Guanajuato. 

Geographic distribution. Central Plateau of Mexico from southern 
Chihuahua to southwestern Mexico (state) and Puebla. (See Fig. 16.) 

Seasonal distribution. June 28 — September 7. 

Records. MEXICO: Chihuahua: Santa Barbara, 6200 ft., September, 9. 
Durango: Durango, 5; Francisco Madero, September, 1; Guadalupe Vic- 
toria, September, 2; Nombre de Dios, July, 1. Guanajuato: State label 



SYSTEMATICS AND PHYLOGENY 



65 



only, 10; 14 mi. south of San Luis de la Paz, September, 6; San Miguel 
Allende, August, 22; Tupataro (Duges, 1889). Hidalgo: Jacala, about 
5000 ft., August, 1; Pachuca, 8000 ft., July, 15. Jalisco: Encarnacion de 
Diaz, July, 2; 12 mi. south of Encarnacion de Diaz, August, 1; 96 km. 
southwest of Guadalajara, 3500 ft., July, 2; La Punta, August, 35. 
Mexico: 20 (highway) mi. southwest of Acambay, 8500 ft., August, 1; 
6 mi. north of Aculco, 6500 ft., August, 3; Atlacomulco, 8100 ft., August, 
3; Temescaltepec, 3. Puebla: Puebla, 1. Queretaro: kilometer 320, near 
Hacienda Balvanera, July, 14. San Luis Potosi: San Luis Potosi, 2. 
Zacatecas: 17 mi. north, 1.5 mi. south, and 9 mi. south of Fresnillo 
(Linsley, MacSwain, and Smith, 1956); Guadalupe, June, 1; 5 mi. south 
of Sombrerete, 7000 ft., September, 1; 2 mi. south of Trancosa, 8000 ft., 
August, 27. 

Remarks. This species shows geographically discordant polymorphic 
variation in the color of the pronotum and the color of the pubescence 
(Fig. 17). The pronotum is either orange with a pair of black discal 
spots or entirely black. In the northern part of the range of the species 
(Chihuahua, Durango, and Zacatecas) the orange phase apparently 




Fig. 17. Polymorphic variation in Lytta variabilis. Circles indicate silver pubescence, 
triangles black, and the square a combination of black and silver. Black area of figures 
indicates percentage of sample with black pronotum and lined area percentage with 
orange pronotum. Number of specimens in each sample is shown by small numerals. 
Where all specimens from a state have the same characters, they are combined to 
form a single sample. 



66 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Table II. Color Phase Composition in Mixed Series of Lytta variabilis 





ORANGE 


BLACK 


TOTAL NUMBER 


LOCALITY 


PHASE 


PHASE 


OF 


SPECIMENS 


Guanajuato (state) 


1 


9 




10 


14 miles south of San Luis de la Paz, 










Guanajuato 


1 


5 




6 


San Miguel Allende, Guanajuato 


16 


6 




22 


Hacienda Balvanera, Queretaro 


3 


11 




14 


Pachuca, Hidalgo 


3 


12 




15 



occurs to the exclusion of the black phase. In the southern part of the 
range mixed series have been taken in Guanajuato, Queretaro, and 
Hidalgo ( Table II ) . I have observed individuals of the two color phases 
feeding on the same plant in Queretaro, and I have seen a male of the 
black phase pinned in copulo with a female of the orange phase from the 
state of Guanajuato. Other southern samples consist of either the black 
or orange phase, but none is large enough to justify the conclusion that 
the population it represents is composed entirely of a single phase. 

Pubescence color varies independently. Over most of the range of the 
species it is silver. In a specimen from Jacala, Hidalgo, and in two speci- 
mens from southwest of Guadalajara, western Jalisco, it is black. In 
three specimens from Temescaltepec, Mexico, it is black on the upper 
surface of the body and on the legs (except coxae) and silver on the 
under surface of the body. 

The specimens from southwest of Guadalajara (a male and a female 
collected by F. G. Werner) have the head and pronotum less coarsely 
punctate than usual for the species, and the sides of the head in the 
male are more deeply concave than in any other male examined. 

The specimens from Temescaltepec (all males) differ from typical 
variabilis in a number of morphological features, as follows: head and 
pronotum much more finely punctate; punctures shallower, only about 
half usual diameter, densely, uniformly arranged but not crowded; 
surface of head and pronotum smooth, shiny, not at all rough or reticu- 
late; pronotal disk more evenly and strongly convex; emargination of 
sixth abdominal sternum more evenly arcuate; lateral lobes of sternum 
more prominent, moderately angulate; and with gonostyli (Fig. 296a) 
relatively short, stubby; aedeagus (Fig. 296b) with dorsal hook more 
massive and ventral hooks closer to each other. In addition, two of the 
specimens have the pygidium (Fig. 205) more strongly produced than 
do any other specimens of variabilis examined. The pygidium is also 
produced in the third specimen but within the normal range of variation 
in variabilis. 



SYSTEMATICS AND PHYLOGENY 67 

The status of the western Jalisco and Temescaltepec populations 
deserves further study. In particular it should be noted that at Toluca, 
Mexico, variabilis occurs in typical form within 50 miles of Temescaltepec. 
I would not be surprised to find that some sort of reproductive isolation 
has been established between typical variabilis and the Temescaltepec 
population, but it seems preferable to assign the Temescaltepec material 
tentatively to variabilis until more information is obtained. 

The type of variabilis has been lost. Following Duges' original inter- 
pretation (that the orange phase is varietal), I have designated as 
neotype a specimen of the black phase. This is a female from "Guana- 
juato" in the British Museum (Natural History). 

Bionomics. With one exception, a specimen from Nombre de Dios, 
Durango, labeled as from Asclepias, all host plants recorded for variabilis 
belong to the family Convolvulaceae. Duges (1869) recorded the type 
material of variabilis from "Convolvulus arbor ea." I collected the speci- 
men from southwest of Acambay, state of Mexico, from a flower of 
Ipomoea stans and the series from south of Trancosa, Zacatecas, from 
near Hacienda Balvanera, Queretaro, and from near San Luis de la Paz, 
Guanajuato, from flowers of I. longifolia. The specimens from southwest 
of Guadalajara, Jalisco, are from "morning-glory flowers." 

Linsley, MacSwain, and Smith ( 1956 ) have recorded variabilis as 
one of a number of insect species competing with the emphorine bee 
Ptilothrix sumichrasti (Cresson) for pollen of Ipomoea longifolia and 
I. pringlei in the Fresnillo region of Zacatecas. These authors also found 
variabilis lightly parasitizing Ptilothrix nesting sites. Whether this para- 
sitism merely reflects the proximity of the bee nests to flowers of 
Ipomoea (P. sumichrasti is oligolectic on Ipomoea) or is in some way 
more directly connected to the host plant relationships of bee and 
parasite is not clear. 

Lytta (Paralytta) scituloides, new species 

Black. Prosternum and gula orange in holotype, black in other speci- 
mens. Pronotum pale orange with a small blackish discal dot on each 
side at middle. Pubescence silver. Wings dark brown. Length: 8 mm.- 
14 mm. 

Differs structurally from variabilis as follows. 

Head with sides nearly parallel above eyes; surface more uniformly 
punctate, as in puberula; punctures not crowded. Pronotum not trans- 
verse, only slightly wider than long; surface smoother, less densely punc- 
tate. 

Male. Head behind eyes deeply, concavely impressed, even more so 
than in scitula. Antennae (Fig. 69) longer, nearly reaching base of 
pronotum; segments proportionately more slender, none wider than long. 



68 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Genitalia with dorsal hook of aedeagus very evenly rounded on dorsal 
margin. 

Female. Antennae nearly as in male except that intermediate segments 
are not enlarged. 

Geographic distribution. Nuevo Leon and eastern Coahuila. (See Fig. 
16.) 

Seasonal distribution. September 13 — March 24. 

Type material. Holotype female and paratype females from [Villa] 
Allende, [Nuevo Leon], October 1917, A. Busck. Allotype male from 
"S J Allende," November 1929. Holotype and allotype in the United 
States National Museum; paratype in my collection. 

Additional records. MEXICO: Coahuila: Saltillo, September, 1. Nuevo 
Leon: Apodaca, March, October, 2; Cola de Caballo [near Monterrey], 
February, 1; Monterrey, October, 2; Villa de Santiago, February, 1. 

Remarks. The head of the male is even more deeply impressed behind 
the eyes in this species than in scitula. However, in scituloides the im- 
pression does not follow the margin of the eye forward, and it therefore 
does not affect the outline of the head in frontal view. The similarity of 
this modification in scituloides and scitula seems to be the result of 
convergence, since other morphological characters indicate that the two 
species are not closely related. Scituloides is actually very similar to 
variabilis and may prove to be only a geographic variant of it. 

Lytta (Paralytta) puberula LeConte 

Lytta puberula LeConte, 1866, Smithsonian Misc. Coll., vol. 6, no. 162, 
p. 162; 1866, Proc. Acad. Nat. Sci. Philadelphia, p. 349. 

Cantharis puberula, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 109. 
Cockerell, 1898, New Mexico Agr. Exp. Sta. Bull. no. 28, p. 173. Fall, 1901, 
Trans. Amer. Ent. Soc, vol. 27, p. 297. Wickham, 1902, Bull. Lab. Nat. Hist. 
State Univ. Iowa, vol. 5, p. 301. 

Black. Wings very pale brown. Pubescence silver, dense, conspicuous 
macroscopically, producing a distinct dusty effect throughout. Length: 
7 mm.-ll mm. 

Differs structurally from variabilis as follows. 

Head usually more uniformly punctate; punctures seldom crowded on 
frontal area and center of vertex, pubescence much longer. Pronotum 
(Fig. 169), as a ride, more evenly rounded at sides, seldom transverse, 
varying from as wide as long to one-tenth wider than long; disk usually 
smoother, more evenly convex; calluses usually small, less distinct, occa- 
sionally obsolete; pubescence fully as long and as conspicuous as on 
vertex. Elytral pubescence much denser, twice as long (as long as or 
longer than that of pronotum ) ; average distance between setae at middle 



SYSTEMATICS AND PHYLOGENY 



69 



of elytra distinctly less than length of a single seta. Tarsal claws (Fig. 
200) more slender and less strongly curved. 

Male. Genitalia as in Figure 298. 

Type locality. Fort Whipple, Arizona. 

Geographic distribution. Northern half of Arizona to extreme south- 
western Colorado and central southern New Mexico. (See Fig. 18.) 

Seasonal distribution. July 17 — September 5. 

Records. UNITED STATES: Arizona: Coconino County, August, 1; 
Flagstaff, July, August, 20; Fort Whipple (LeConte, 1866); Grand Can- 
yon, July, 1; 5 mi. south of Grand Canyon, 6800 ft., August, 1; McNary, 
August, 4; Oak Creek Canyon, 6000 ft., 1; Peach Springs, Hualpai Indian 
Reservation, September, 2; Prescott, 12; Show Low, August, 2; White 
Mountains, August, 2; Williams, July, August, 9; Yavapai County, July, 
August, 28. Colorado: Durango, July, 5; Mesa Verde, July, 1. New 
Mexico: Cloudcroft, 9000 ft., 7 (4 in July); Continental] D[ivide,] 
Ten[aja] (not located), 2; Fort Wingate, 1. 

Remarks. The distal antennal segments in ptiberula tend to be a little 
more slender proportionately than in variabilis, but at best this difference 
is a very subtle one. 




Fig. 18. Distribution of Lijtta puberula. 



70 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

It is unlikely that any difficulty will be encountered in recognizing 
this species since the only other member of the Variabilis Group occur- 
ring in the United States is biguttata. Moreover, the dusty effect produced 
by its dense pubescence separates puberula from all other black forms 
of Lytta except one of the color varieties of the Mexican species proteus. 

This species is unusual in two respects. It is the only species of the 
subgenus Paralytta known to occur on plants of the family Campanu- 
laceae, and its range is disjunct in relation to the ranges of the other 
members of its subgroup. According to my interpretation of relationships, 
puberula is a very close and somewhat specialized relative of variabilis. 
Evidently, at one time the range of the common ancestor of these two 
species extended north well into Arizona and New Mexico. Subsequently, 
its range was interrupted through the lower regions of these states and 
adjacent Mexico, with the puberula line persisting as a population adapted 
to conditions at higher elevations in the north and the variabilis line 
being restricted to the central and southern portions of the Central 
Plateau. 

The type of puberula is in the Museum of Comparative Zoology. 

Bionomics. Four specimens are recorded from Campanula (Campanu- 
laceae) at Cloudcroft, New Mexico. 

Lytta (Paralytta) michoacanae (Champion) 

Cantharis michoacanae Champion, 1892, Biol. Centrali-Americana, Coleop- 
tera, vol. 4, pt. 2, p. 447. 

Lytta michoacanae, Selander, 1954, Jour. Kansas Ent. Soc, vol. 27, p. 89. 

Black (a little brassy in fresh material). Prothorax orange; pronotum 
with a black discal spot on each side at middle. Wings pale brown. 
Pubescence black on upper surface, silver on under surface, dark but 
with a golden luster on legs. Length: 7 mm. -11 mm. 

Head with sides parallel or convergent above eyes; surface smooth, 
very shiny, finely, very sparsely punctate; pubescence very short, erect. 
Eyes unusually large, very prominent. Pronotum subcircular, as wide as 
or barely wider than long; disk transversely, shallowly impressed before 
middle, evenly convex behind, not impressed on midline or at base; discal 
spots elevated on impunctate calluses; basal margin more deeply emar- 
ginate than in variabilis; surface finely, very sparsely punctate at sides, 
nearly impunctate at center; pubescence consisting of a few very short, 
erect setae. Elytra rugose, lacking distinct punctures except at base; 
glabrous (actually with a few very short setae visible in proper light). 
Tarsi more slender than in related species. 

Male. Head behind eyes clothed with extremely short setae but other- 
wise unmodified. Antennae (Fig. 70) reaching base of pronotum; seg- 
ments IV and V subequal in size, roughly as wide as VI, a little 



SYSTEMATICS AND PHYLOGENY 



71 



compressed but not perceptibly incrassate except in comparison with 
same segments in female; VI-VIII progressively slightly wider; segments 
longer than broad. Sixth abdominal sternum as in Figure 226; lateral lobes 
of sternum obtuse. Pygidium not produced. Genitalia as in Figure 299. 

Female. Antennae as in male except that segments IV and V are visibly 
narrower than VI. 

Type locality. Huetamo, Michoacan. 

Geographic distribution. Lower elevations from central Michoacan to 
eastern Guerrero. (See Fig. 19.) All known localities for michoacanae 
are within the valley system of the Rio Balsas and its tributaries. 

Seasonal distribution. July 13 — August 20. 

Records. MEXICO: Guerrero: Iguala, 3; 30 km. south of Iguala, 
2300 ft., farmland, July, 25; Mexcala [Mezcala], 1350 ft., August, 1; 5 mi. 
north-northeast of Mexcala, 2000 ft., August, 5; 9 km. north of Zumpango 
[del Rio], 2800 ft., August, 18. Michoacan: Huetamo, 2 (paratypes); 
Tacambaro [de Codallos] (Champion, 1892). 

Remarks. Specimens of this species superficially resemble small speci- 
mens of the orange phase of variabilis but are easily separable on the 
basis of the color of the pubescence on the upper surface of the body 
(pale in orange-phase variabilis) as well as by structural characters. 




Fig. 19. Distribution of Lytta michoacanae (circles) and Lijtta ebenina ( squares ] 



72 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

The material from Guerrero agrees in every detail with the two para- 
types I have seen from Michoacan. The type is in the British Museum 
(Natural History). 

Bionomics. The specimens from 9 km. north of Zumpango del Rio and 
from 30 km. south of Iguala, Guerrero, were collected on flowers of 
Convolvulaceae. 

Lytta (Paralytta) ebenina (Duges) 

Cantharis ebenina Duges, 1877, La Naturaleza, vol. 4, p. 66, pi. 2, fig. 10; 
1889, An. Mus. Michoacano, vol. 2, p. 103. Champion, 1892, Biol. Centrali- 
Americana, Coleoptera, vol. 4, pt. 2, p. 443. 

Black. Wings pale brown. Pubescence black. Length: 10 mm. -15 mm. 

Head transverse, decidedly wider than long, strongly triangular; tem- 
pora prominent; surface finely granular, dull; vertex finely, sparsely 
punctate; frontal area more densely punctate, but punctures not crowded; 
pubescence short, erect. Pronotum transversely suboval, at least one-tenth 
wider than long; disk flattened, with a large impunctate area on each 
side but without distinct calluses; surface as on vertex but more irregu- 
larly punctate; center of disk nearly glabrous. Elytra obsolescently 
rugose, nearly smooth, shinier than head and pronotum, impunctate, 
nearly glabrous (with only a few very short setae). Tarsal claws heavy, 
strongly curved. 

Male. Head behind eyes flattened. Antennae (Fig. 71) not quite 
reaching base of pronotum; segments 1II-VI weakly incrassate; IV larg- 
est, as wide as or a little wider than X; V and VI slightly smaller, both 
wider than VII; segments longer than broad. Sixth abdominal sternum 
as in Figure 227. Pygidium not produced. Genitalia as in Figure 300. 

Female. Antennae (Fig. 122) reaching just beyond middle of pronotum. 
Pad of first segment of middle tarsi limited to apical half, of hind tarsi 
absent. 

Type locality. Mexico. 

Geographic distribution. Tlaxcala and southern Puebla to Tabasco 
(La Venta ruins). (See Fig. 19.) 

Seasonal distribution. May — June 29. 

Records. MEXICO: Country label only, 6. Puebla: Amozoc, July, 13; 
Esperanza, 5; Matamoros Iziicar [Izucar de Matamoros] (Champion, 
1892). Tabasco: La Venta [ruins], May, 4. Tlaxcala: 4 mi. and 5 mi. east 
of Apizaco, June, 342; highway 18, border of Tlaxcala and Puebla, July, 
1; 4 mi. northwest of Huamantla, June, 31; 2.5 mi. west of El Carmen, 
June, 27. Veracruz: Boca del Monte (Duges, 1889); San Andres [Tuxtla], 
1. 



SYSTEMATICS AND PHYLOGENY 73 

Remarks. This species is distinguished from all other black species 
of Lytta by the combination of dull, finely granular head and pronotum; 
shiny, nearly smooth elytra; and the absence of a pale frontal spot. 

There is a specimen in the Haag-Rutenberg collection in the Zoologis- 
che Staatsammlung in Munich labeled as from Panama (Koltze), but 
this locality is almost certainly in error. 

The type of ebenina has been lost. As neotype I have designated a 
male from San Andres Tuxtla, Veracruz, in the British Museum (Natural 
History ) . 

Bionomics. The three series of specimens recorded from the state of 
Tlaxcala were collected in 1955 from the flowers of Argemone platyceras 
( Papaveraceae ) . Feeding seemed to be confined to pollen. At each of 
the localities ebenina was associated on the host plant with Li/tta quadri- 
macnlata. 

Koltzei Subgroup 

Black or dark metallic blue or green throughout, or with pronotum 
pale, bimaculate. Head lacking a pale frontal spot. First segment of 
male middle tarsi unmodified or slightly thickened and weakly bent. 
Male sixth abdominal sternum deeply emarginate; lateral lobes of sternum 
produced. Female sixth sternum shallowly, triangularly emarginate at 
apex. 

Remarks. This subgroup contains the species plumbea, koltzei, canelas, 
augusti, and bipuncticollis. Except for canelas, which has been recorded 
only from a single locality in Durango, the species of the subgroup are 
restricted in range to southern Mexico. Based on records for only two 
species (koltzei and plumbea), the host plants of the subgroup are 
species of Convolvulaceae and Papaveraceae. 

Key to Species 

1. Antennae longer, reaching or extending beyond base of pronotum in 
male, reaching at least basal third of pronotum in female; pronotum 
strongly alutaceous, coarsely, densely punctate; male middle tarsi 
unmodified; lateral lobes of male sixth abdominal sternum not appre- 
ciably divergent; aedeagus with both ventral hooks well developed; 

pronotum black 2 

Antennae, shorter, not extending beyond basal third of pronotum in 
male, not reaching middle of pronotum in female; pronotum rela- 
tively smooth, less coarsely, less densely punctate; first segment of 
male middle tarsi slightly thickened, weakly bent; lateral lobes of 
male sixth abdominal sternum divergent; aedeagus with apical ventral 
hook small or obsolete 4 



74 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

2. Emargination of male sixth abdominal sternum broadly triangular, 
extending at most one-third length of sternum (Fig. 228); lateral 
lobes of sternum broadly triangular; elytra black with a tinge of blue, 
not costate, essentially glabrous; pubescence pale on under surface, 

black elsewhere plumbea (p. 74 ) 

Emargination of male sixth abdominal sternum U-shaped, extending 
at least half length of sternum ( Figs. 229-30 ) ; lateral lobes of sternum 
slender 3 

3. Elytra dark blue or green, not costate, moderately densely, conspicu- 
ously pubescent; pubescence silver on at least elytra and under sur- 
face. Michoacan to Puebla koltzei (p. 76) 

Elytra black, tricostate, essentially glabrous; pubescence black 
throughout. Durango canelas (p. 79) 

4. Pronotum black; aedeagus with apical hook small (Fig. 304c) 

augusti (p. 80 ) 

Pronotum orange; aedeagus with apical hook obsolete (Fig. 305c) . . . 
bipancticollis (p. 82 ) 

Lytta (Paralytta) plumbea Haag-Rutenberg 

Cantharis plumbea Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 
24, p. 38. 

Cantharis monilicornis Duges, 1881, La Naturaleza, vol. 5, p. 142, pi. 4, 
fig. 3; 1889, An. Mus. Michoacano, vol. 2, p. 96. New synonymy. 

Cantharis koltzei, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 444. Misidentification. 

Black tinged with dark blue. Upper surface, particularly head and 
pronotum, with a brassy or iridescent luster, often faint. Wings brown. 
Pubescence golden or silver on under surface, black elsewhere. Length: 
7 mm. -14 mm. 

Head with tempora much more well defined and quadrate than in 
koltzei; surface canaliculate, shiny, coarsely, deeply, very densely punc- 
tate; punctures crowded on frontal area and most of vertex; pubescence 
moderately long, erect. Eyes smaller and less prominent than in koltzei; 
head at level of eyes at most only slightly wider than at tempora, usually 
not as wide. Pronotum as in Figure 170, as wide as or a little wider 
than long; disk uneven, transversely impressed just before middle; a 
deep, groovelike impression on midline from near middle to base; calluses 
impunctate, often small, poorly defined; two shallow foveae often present 
on each side of disk, one near basal corner and another just behind callus; 
basal margin of pronotum strongly recurved on midline; surface roughly 
alutaceous, shiny, coarsely, densely punctate; pubescence as on vertex. 
Elytra rugose punctate basally, merely rugose elsewhere, essentially 
glabrous, with a few setae at base and apex. 



SYSTEMATICS AND PHYLOGENY 



75 



Male. Head behind eyes not noticeably flattened. Antennae (Fig. 72) 
reaching base of pronotum; segments IV-VI moderately incrassate; IV 
largest, V and VI progressively smaller; VI barely wider than X; VII-X 
more moniliform than preceding segments, progressively barely wider, 
not much longer than wide. First segment of middle tarsi unmodified. 
Fifth abdominal sternum feebly emarginate, almost entire. Emargination 
of sixth sternum (Fig. 228) deeply, broadly subtriangular, extending 
one-fifth to one-third length of sternum; lateral lobes of sternum moder- 
ately produced, broadly triangular. Pygidium weakly produced. Genitalia 
as in Figure 301, very similar to those of koltzei; aedeagus with two 
well-developed ventral hooks, these more widely separated than in re- 
lated species. 

Female. Antennae (Fig. 123) reaching about basal one-third of pro- 
notum. Sixth abdominal sternum (Fig. 275) very shallowly emarginate 
at apex. 

Type locality. Of plumbea, Mexico; of monilicornis, Uruapan [del 
Progreso], Michoacan. 

Geographic distribution. Central Plateau of Mexico from Guanajuato 
and nothern Michoacan to the Distrito Federal. (See Fig. 20.) 




Fig. 20. Distribution of Lytta plumbea (circles) and Lytta augusti (squares). 



76 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ll/tta 

Seasonal distribution. June 26 — August 14. 

Records. MEXICO: Country label only, 1 (type of plumbea). Distrito 
Federal: Mexico City, 1. Guanajuato: Guanajuato, 1; Moroleon (Duges, 
1889). Mexico: Toluca, 8800 ft., July, August, 37; 1.8 mi. southeast of 
Toluca, 8800 ft., July, 406. Michoacan: Carapan, July, 1; Patzcuaro, 
6500 ft., August, 2; Quiroga (Duges, 1889); Tancitaro, 6000 ft.-6600 ft., 
August, 29; Uruapan [del Progreso] (Duges, 1881). 

Remarks. Lytta plumbea resembles koltzei but is distinguished in both 
sexes by its darker, less metallic coloration, heavier and shorter antennae, 
and subglabrous elytra, as well as by differences in the shape of the 
pronotum and the sixth abdominal sternum. 

Some variation in details of surface sculpturing and in the relative 
lengths of the antennal segments is evident in the long series of speci- 
mens studied. Otherwise the species seems to be relatively stable in its 
structural characters. 

Champion thought Duges ( 1889, p. 97 ) referred to koltzei ( cyanescens ) 
in recording a metallic variety of plumbea (monilicornis) from Quiroga, 
Michoacan, but this is not a safe assumption in view of the range of 
color variation in plumbea. 

As lectotype of plumbea I have designated a female in the Zoologische 
Staatssammlung in Munich ( Haag-Rutenberg collection ) labeled "Mexico, 
Klug, Coll. Germar." 

Duges' type apparently having been lost, I have designated as neotype 
of monilicornis a male in the British Museum (Natural History) from 
Guanajuato, [Guanajuato] (Salle collection). 

Rionomics. The specimens from Toluca and vicinity, state of Mexico, 
and the specimen from Carapan, Michoacan, were collected on Argemone 
platyceras ( Papaveraceae ) . Four specimens from Tancitaro, Michoacan, 
labeled as from poppy, are probably from Argemone also. In the Toluca 
area the beetles occurred in the flowers of the host plant either singly or 
in groups of two to six. Some feeding on the petals was observed, but 
it was concluded that the principal food material of the adults is pollen. 
One or more individuals of Lytta quadrimaculata were present in about 
half the flowers containing plumbea. 

In addition to the above data, there is a record of two specimens of 
plumbea on flowers of morning-glory ( Convolvulaceae ) at Patzcuaro, 
Michoacan. 

Lytta (Paralytta) koltzei Haag-Rutenberg 

Lytta koltzei Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 24, p. 38. 
Lytta koltzei var. cyanescens Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., 
vol. 24, p. 38. New synonymy. 



SYSTEMATICS AND PHYLOGENY 77 

Lytta humilis Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 24, p. 39. 
Denier, 1940, Rev. Ent., vol. 11, p. 802. New synonymy. 

? Cantharis atrovirens Duges, 1889, An. Mus. Michoacano, vol. 2, p. 104. 
Champion, 1892, Biol. Centrali-Americana, Coleoptera, vol. 4, pt. 2, p. 449. 
New synonymy. 

Cantharis cyanescens, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 444. 

Cantharis humilis, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 445. 

Lytta koltzei var. minor Borchmann, 1917, Coleopterorum catalogus, pt. 
69, p. 95. 

Head and pronotum black with an iridescent luster, reflecting blue, 
violet, and green. Elytra metallic dark blue or bluish green. Under sur- 
face and femora black with a distinct metallic dark blue luster. Tibiae 
and tarsi black. Wings brown. Pubescence silver, piceous tipped with 
silver, or entirely piceous on head and pronotum; silver on elytra, under 
surface, and femora; usually piceous on tibiae and tarsi. Length: 7 mm.- 
13 mm. 

Head subcanaliculate, shiny, coarsely, deeply, densely punctate; punc- 
tures crowded on frontal area and center of vertex; pubescence moder- 
ately long, erect. Eyes large; head clearly wider at level of eyes than 
at tempora. Pronotum as in Figure 171, varying from one-tenth longer 
than wide to slightly wider than long; disk uneven, transversely impressed 
just before middle and on midline at base; calluses sometimes indistinct, 
never strongly elevated; surface roughly alutaceous, shiny, coarsely, 
densely punctate; pubescence as on vertex. Elytra rugose punctate 
basally, merely rugose elsewhere; pubescence moderately long, semi- 
erect, moderately dense, conspicuous; average distance between setae 
equal to length of a single seta. 

Male. Head behind eyes not noticeably flattened. Antennae (Fig. 73) 
reaching one or two segments beyond base of pronotum; segments 
IV-VI weakly incrassate; IV and V subequal in size; VI slightly narrower 
than V, barely wider than X; VII like VIII or intermediate in size and 
form between VI and VIII; VIII-X more moniliform than preceding 
segments; segments elongate, IV-X at least one-half longer than wide. 
First segment of middle tarsi unmodified. Fifth abdominal sternum 
moderately deeply emarginate at center. Emargination of sixth sternum 
(Fig. 229) very deep, U-shaped, extending more than half length of 
sternum; lateral lobes of sternum strongly produced as slender, nearly 
parallel processes, either subtruncate or rounded at apex. Pygidium not 
produced. Genitalia as in Figure 302 ( gonostyli in lateral view sometimes 
more bluntly rounded at apex than shown); aedeagus with two well- 
developed hooks. 



78 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ll/ttCl 



Female. Antennae (Fig. 124) reaching base of pronotum; distal seg- 
ments less elongate than in male, much as in female of plumbea and cane- 
las. Sixth abdominal sternum as in Figure 276 or more shallowly 
emarginate than shown and with blunter lateral lobes (as in canelas). 

Type locality. Of cyanescens and atrovirens, Mexico. The type locality 
of koltzei and of humilis is supposedly Panama, but this is probably in- 
correct (see below). 

Geographic distribution. Central Michoacan to central eastern Puebla. 
(See Fig. 21.) 

Seasonal distribution. August 3 — October 24. 

Records. MEXICO: Country label only, 4 (including a paratype of 
koltzei). Distrito Federal: Mexico City, 1; Tlalpam [Tlalpan], September, 
1. Mexico: Real de Arriba, [near] Temescaltepec, 1; Temescaltepec, 19. 
Michoacan: Pdtzcuaro, 6500 ft., August, 2. Morelos: 6 mi. north of Cuer- 
navaca, 7500 ft., August, 3; 9 km. south of Tres Cumbres, 7700 ft., August, 
6; Tres Marias [Tres Cumbres], October, 1. Puebla: Esperanza, 1. 

PANAMA [?]: Country label only, 5 (from type series of koltzei and 
humilis). 

Remarks. In addition to the male sexual characters, the conspicuously 




Fig. 21. Distribution of Lijtta koltzei (circles) and Lytta canelas (square). 



SYSTEMATICS AND PHYLOGENY 79 

pubescent elytra readily distinguish this species from other species of 
the Koltzei Subgroup. Structurally it most closely resembles canelas. 

The type series of koltzei consists of three females. Two of these 
specimens (including one which I have designated as lectotype) are 
labeled "Panama, Koltze;" the other specimen is labeled "Mexico, Bates." 
As lectotype of the variety cyanescens I have designated a male labeled 
"Mexico, V. Hyd." The type series of humiUs consists of two males 
and a female labeled with the same data as the type of koltzei, and in 
the same hand; I have designated as lectotype one of the males. I have 
carefully compared all the material mentioned above and am quite cer- 
tain that only a single species is represented. All of it is in the Zoologische 
Staatssammlung in Munich. 

The label "Panama, Koltze" mentioned above would seem to be 
erroneous inasmuch as there have been no other records of the present 
species south of the state of Puebla, Mexico. In this connection it should 
be stated that as far as I know, the only other specimen of Lytta in the 
Haag-Rutenberg collection bearing this particular label is a representative 
of ebenina, another species which has not otherwise been recorded south 
of Mexico ( the state of Tlaxcala in this case ) . 

Cantharis atrovirens was described by Duges from a single specimen, 
since lost. Its description could apply to koltzei except that the outer hind 
tibial spur is said to have the "form of a spear." I have placed atrovirens 
as a questionable synonym of koltzei on the assumption that either Duges' 
description is inexact or that the specimen he described was aberrant. 

Bionomics. The specimens from near Tres Cumbres, Morelos, were 
collected on flowers of morning-glory ( Convolvulaceae ) . 

Lytta (Paralytta) canelas, new species 

Cantharis plumbea, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 443. Misidentification. 

Lytta plumbea, Vaurie, 1950, Amer. Mus. Novitates, no. 1477, p. 43. Mis- 
identification. 

Black. Head and pronotum with a faint brassy luster. Wings brown. 
Pubescence black. Length: 9 mm. -12 mm. 

Head somewhat canaliculate, moderately shiny, coarsely, deeply, 
densely punctate; punctures for most part crowded; pubescence moder- 
ately long, erect. Pronotum as in Figure 172 or more nearly circular than 
shown, barely wider than long; disk with a deep transverse impression 
just before middle and an equally deep one on midline at base; basal 
margin strongly recurved; calluses strongly elevated, impunctate; a large, 
distinct fovea just behind each callus, sometimes joined with basal im- 
pression; surface roughly alutaceous, moderately shiny, coarsely, densely 
punctate; pubescence as on vertex. Elytra unique for Lytta in being 



80 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

distinctly costate; three costae present on each elytron, producing an 
undulate surface; first costa extending from base midway between scu- 
tellum and humerus to near apex, separated from second costa by a 
depression containing a nervule; second costa extending from humerus 
almost to apex; third costa less well defined than others, located on in- 
flexed part of elytron, fused with second costa at humerus and not extend- 
ing much beyond middle of elytron; surface of elytra rather dull, rugose 
punctate except near apex (where punctures become indistinct), essen- 
tially glabrous, with only a few very short setae. 

Male. Head behind eyes flattened, clothed with setae about half as 
long as those on frontal area. Antennae reaching base of pronotum; 
segments IV-VI weakly incrassate; IV largest, a little wider than X; 
V and VI progressively slightly smaller, both wider than VII; VII-X 
more moniliform than preceding segments, progressively barely wider; 
segments longer than wide. First segment of middle tarsi unmodified. 
Fifth abdominal sternum shallowy emarginate. Emargination of sixth 
sternum (Fig. 230) very deep, U-shaped, extending half length of ster- 
num; lateral lobes of sternum strongly produced as slender, parallel 
processes, rounded at apex. Pygidium not produced. Genitalia as in 
Figure 303; aedeagus with two well-developed ventral hooks. 

Female. Antennae not reaching base of pronotum. Sixth abdominal 
sternum as in Figure 277, shallowly emarginate at apex. 

Geographic distribution. Known only from the type locality, in west- 
ern Durango. (See Fig. 21.) 

Seasonal distribution. Unknown. 

Type material. Holotype male, allotype female, and three paratypes 
(one male, two females) from Canelas, Durango, collected by Becker. 

Remarks. The costate elytra immediately distinguish canelas from 
all other species of Lytta. Structurally canelas is most similar to koltzei. 

The present description is based on five of the six specimens which 
Champion mistakenly assigned to plumbed. I have not seen the sixth 
specimen of his series. 

Lytta (Paralytta) augusti Haag-Rutenberg 

Lytta variabilis var. augusti Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., 
vol. 24, p. 38 (footnote). 

Cantharis subviolacea Champion, 1892-93, Biol. Centrali-Americana, Cole- 
optera, vol. 4, pt. 2, pp. 445, 463, pi. 21, fig. 22. 

Lytta augusti, Selander, 1954, Jour. Kansas Ent. Soc., vol. 27, p. 89, fig. 1. 

Head and pronotum black with a brassy luster. Elytra metallic dark 
blue or purple, or black with a blue or purple luster. Under surface of 
abdomen and femora black with a metallic blue luster. Under surface 



SYSTEM ATICS AND PHYLOGENY 81 

of thorax, tibiae, and tarsi black. Wings pale brown. Pubescence fine, 
either pale with black base or entirely pale. Length: 11 mm. -15 mm. 

Head shiny, moderately densely, coarsely to finely punctate, with a 
tendency when coarsely punctate for partial fusion of punctures, produc- 
ing short canaliculations; frontal area impunctate and impressed at center 
between eyes; pubescence moderately long, erect. Pronotum suboval, a 
little more than one-tenth wider than long; disk rather even, somewhat 
flattened before middle; calluses at least weakly elevated; a fovea usually 
present just behind each callus; midline impunctate, impressed, impres- 
sion widening at base; surface not alutaceous, very shiny; punctures as 
on vertex or much sparser, especially at center; pubescence slightly 
shorter than on vertex. Elytra rugose, lacking distinct punctures except 
for some fine ones near base; pubescence short, semierect, very sparse; 
setae inconspicuous unless entirely pale, separated by a distance as 
great as or greater than length of a single seta. Outer hind tibial spur 
spoon-shaped. 

Male. Head behind eyes flattened. Antennae (Fig. 74) short, at most 
reaching basal third of pronotum; segments III-V weakly incrassate; III 
barely wider than in female; IV largest, as wide as VI; V slightly en- 
larged in comparison with same segment in female; VI-X progressively 
slightly wider; segments short, at least in part wider than long. Middle 
tibiae nearly straight. First segment of middle tarsi very slightly thick- 
ened, very weakly bent in dorsal view (as in bipitncticollis) . Fifth ab- 
dominal sternum shallowly emarginate. Emargination of sixth sternum 
(Fig. 231) very deep, somewhat triangular, extending half length of 
sternum; lateral lobes of sternum strongly produced as slender, moder- 
ately divergent processes, rounded at apex. Pygidium not produced. 
Genitalia as in Figure 304; aedeagus with two ventral hooks, but apical 
hook extremely small. 

Female. Antennae (Fig. 126) not reaching middle of pronotum; seg- 
ments III and IV equal in size. Sixth abdominal sternum with a small 
triangular emargination at apex; lateral lobes of sternum obtuse. 

Type locality. Of augusti, Mexico; of subviolacea, [La] Parada or 
Durasnal, both in Oaxaca. 

Geographic distribution. Recorded from central Oaxaca and the Vera- 
cruz-Puebla border southwest of Orizaba. (See Fig. 20.) 

Seasonal distribution. The only recorded collection date is August 1. 

Records. MEXICO: Country label only, 2 (including lectotype). 
Oaxaca: Durasnal (not located), 1 (paratype of subviolacea); [La] 
Parada, 1 (paratype of subviolacea); Oajaca [Oaxaca de Juarez], 2. 
Veracruz: El Puerto, 7000 ft., 31 mi. southwest of Orizaba, August, 2. 



82 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Remarks. In a male and female from El Puerto, Veracruz, all clothing 
setae are entirely silver. In the other specimens examined the setae of 
the upper surface are piceous tipped with gold or silver, while the setae 
of the under surface are silver for most of their length, becoming piceous 
at the immediate base. 

The specimen (a female) from La Parada, Oaxaca, exhibits bilaterally 
symmetrical distortion of the fore and middle legs. The femur and tibia 
of one of the middle legs of this specimen are shown in Figure 193. The 
fore legs are similarly distorted, except that the femora and tibiae are not 
bent. 

The Haag-Rutenberg series of variabilis in the Zoologische Staatssamm- 
lung in Munich includes a male and two females of augusti, all bearing 
the label "Mexico, Deyrolle." These are unquestionably the specimens 
referred to as "variabilis var. augusti" by Haag-Rutenberg. I have desig- 
nated one of them (the male) as the lectotype of augusti. The type of 
subviolacea is in the British Museum (Natural History). 

Lytta (Paralytta) bipuncticollis Haag-Rutenberg 

Lytta bipuncticollis Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 
24, p. 36. 

Cantharis bipuncticollis, Champion, 1892, Biol. Centrali-Americana, Cole- 
optera, vol. 4, pt. 2, p. 443, pi. 20, fig. 21. 

Head black. Pronotum orange, finely margined with black, and with 
a black discal spot on each side at middle. Elytra metallic dark bluish 
purple or green. Under surface and femora black with a metallic blue 
or bluish green luster. Tibiae and tarsi black. Wings brown. Pubescence 
black. Length: 12 mm.-17 mm. 

Head smooth, shiny, finely, sparsely, irregularly punctate; an impunc- 
tate area at center of frontal area between eyes; pubescence moderately 
long, erect. Pronotum suboval, varying from slightly wider than long 
to fully one-fifth wider than long; sides more strongly convergent apically 
than basally; disk even, somewhat flattened, especially at center before 
middle; midline impunctate; impressed; discal spots elevated on im- 
punctate calluses; surface smooth, finely, sparsely punctate; pubescence 
short, sparse, erect. Elytra rugose, lacking distinct punctures except for 
some fine ones near base; pubescence fine, sparse, semierect, difficult to 
detect except in proper light; setae at middle of elytra separated by an 
average distance about equal to length of a single seta. Outer hind tibial 
spur spoon-shaped. 

Male. Head behind eyes flattened. Antennae short, reaching middle 
of pronotum; segments III-VI incrassate; III barely wider than in female; 
VI wider than VII but similar in form; VII-X progressively slightly 
wider; segments short, VI-X as wide as or wider than long. In one male 



SYSTEMATICS AND PHYLOGENY 



83 



(Cordoba, Veracruz) segments IV and V are unusually large, IV being 
wider than, and V as wide as, X; in the other male studied (Mochitlan, 
Guerrero ), IV is not wider than V and both are only about as wide as VII. 
Middle tibiae a little expanded in comparison with those of female. 
First segment of middle tarsi slightly thickened, very weakly bent in 
dorsal view. Fifth abdominal sternum shallowly emarginate. Emargina- 
tion of sixth sternum (Fig. 232) very deep, triangular, extending less 
than half length of sternum; lateral lobes of sternum strongly produced 
as slender, strongly divergent processes, subacute at apex. Pygidium not 
produced. Genitalia as in Figure 305; aedeagus with a single ventral hook 
(apical hook obsolete). 

Female. Antennae not reaching middle of pronotum; segment III 
slightly wider than IV. Sixth abdominal sternum as in Figure 278; lateral 
lobes of sternum acute. 

Type locality. Mexico. 

Geographic distribution. Central Veracruz (Jalapa) to eastern Guer- 
rero and central Oaxaca. ( See Fig. 22. ) 

Seasonal distribution. Unknown. 

Records. MEXICO: Country label only, 4 (including lectotype). Guer- 
rero: State label only, 4; Amula (Champion, 1892); Mochitlan, 1. Oaxaca: 




Fig. 22. Distribution of Lytta bipuiicticollis. 



84 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LljttCl 

Oaxaca [de Juarez] (Champion, 1892). Veracruz: Cordoba, 2; Jalapa 
(Champion, 1892). 

Remarks. From all other North American species of Lytta having an 
orange or red pronotum, bipuncticollis is easily recognized by its bluish 
or greenish elytra. 

Structurally bipuncticollis most closely approaches augusti. In males 
of both species the pygidium may overlap the sixth abdominal sternum 
to the extent that the true shape of the latter is obscured. 

Included among the material examined are 3 of the 15 specimens 
studied by Champion for the Biologia Centrali-Americana report. As 
lectotype of bipuncticollis I have designated a male labeled "Mexico, V. 
Heyd." in the Zoologische Staatssammlung in Munich ( Haag-Rutenberg 
collection ) . 

Quadrimaculata Subgroup 

Head when dark with a pale frontal spot. Pronotum usually pale, 
bimaculate, varying occasionally to entirely black. Elytra partially or 
entirely pale (except some proteus and scitula). Under surface and 
legs often pale, at least in part. First segment of male middle tarsi 
bent in dorsal view, excavate on anterior side for reception of anterior 
middle tibial spur (except in sonorae). Male sixth abdominal sternum 
deeply emarginate; lateral lobes of sternum produced. Female sixth ster- 
num truncate (except in biguttata). Pygidium produced. 

Remarks. This subgroup contains the species sonorae, biguttata, sci- 
tula, quadrimaculata, and proteus. Except for biguttata, the species of 
the subgroup are not known to occur outside Mexico. However, it is 
not unlikely that sonorae, now recorded from Sonora, ranges north into 
Arizona. Scitula and proteus seem to be restricted to southern Mexico. 
Quadrimaculata is widely distributed on and south of the Central Plateau. 
Riguttata is the most widely distributed species of Lytta in North 
America, with a range extending from Montana to near the southern end 
of the Central Plateau of Mexico. Adults of the Quadrimaculata Sub- 
group feed on species of Convolvulaceae, Papaveraceae, and Compositae. 

All the species of the subgroup except sonorae are extremely variable 
in color in the adult stage. This is in contrast to the condition in the 
species of the Variabilis and Koltzei subgroups where color is stable or 
relatively so ( the color of the pronotum and pubescence is geographically 
variable in variabilis) . 

Key to Species 

1. Emargination of male sixth abdominal sternum at least as wide as 
deep, clearly not dividing visible portion, extending less than half 



SYSTEMATICS AND PHYLOGENY 85 

actual length of sternum; male gonostyli neither extremely broad nor 

truncate; aedeagus with a pair of ventral hooks 2 

Emargination of male sixth abdominal sternum much deeper than 
wide, nearly or completely dividing visible portion, extending nearly 
three-fourths actual length of sternum (Fig. 174); male gonostyli 
extremely broad in lateral view, truncate (Fig. 309b); aedeagus with 
a single ventral hook (Fig. 309c) proteus (p. 97) 

2. Male middle tarsi not modified; intermediate segments of male anten- 
nae weakly incrassate (Fig. 75); elytra each with a long, dark discal 

vitta reaching or approaching humerus and apex. Sonora 

sonorae (p. 85 ) 

First segment of male middle tarsi bent, excavate basally on anterior 
side; intermediate segments of male antennae at least in part strongly 
incrassate; elytra not as above 3 

3. Male pygidium strongly, narrowly prolonged medially (Fig. 209); 
elytra orange, each usually with a black spot at apical and basal third 

( Fig. 159 ) , rarely with one or both spots absent 

quadrimaculata (p. 93) 

Male pygidium not strongly, narrowly prolonged medially; elytra 
never with a black spot at basal third 4 

4. Male head behind eyes slightly flattened, not concave; elytra yellow, 
usually with a black spot, short vitta, or pair of vittae on each at 

apical fourth (Fig. 160). United States and Mexico 

biguttata (p. 86) 

Male head behind (and above) eyes deeply concave; elytra black, 
each often with lateral margin and a broad discal vitta yellow (Fig. 
161 ). Southern Mexico scitula (p. 90) 

Lijtta ( Paralytta ) sonorae Van Dyke 
Lytta sonorae Van Dyke, 1947, Pan-Pacific Ent, vol. 23, p. 156. 

Orange. Antennae, palpi, and tarsi brown. Pronotum with a small black 
discal spot on each side just behind middle and a smaller spot on lateral 
margin of disk just before middle. Under surface of thorax sometimes 
partially suffused with black. Each elytron with a broad brown discal 
vitta which begins at or near humerus and which may or may not reach 
apex. Wings colorless. Pubescence pale golden or silver. Length: 7.5 mm.- 
8.0 mm. 

Head with surface smooth, shiny, rather finely, sparsely punctate; 
pubescence scattered, erect. Pronotum as in biguttata, but basal margin 
more shallowly emarginate at center. Elytral surface as in quadrimacu- 



86 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

lata, more strongly rugose than in biguttata; pubescence conspicuous, 
regularly distributed, semierect. 

Male. Head behind eyes weakly flattened. Antennae (Fig. 75) reach- 
ing base of pronotum; segments III-VI weakly incrassate; IV largest, as 
wide as X, not curved; VII-X a little longer than wide. First segment 
of middle tarsi unmodified. Fifth abdominal sternum entire. Emargina- 
tion of sixth sternum (Fig. 233) deeply semicircular; sternum impressed 
at center; lateral lobes of sternum relatively broad, subacute. Pygidium 
produced as in Figure 206; apex acute. Genitalia similar in most respects 
to those of biguttata; aedeagus as in Figure 297. 

Female. Sixth abdominal sternum truncate at apex. 

Type locality. Carbo, Sonora. 

Geographic distribution. This is the only species of the Variabilis 
Group except biguttata known to occur in the Sonoran Desert. It has been 
recorded only from the type locality, which is approximately 45 miles 
north of Ffermosillo, Sonora. (See Fig. 23.) 

Seasonal distribution. Collected November 24. A relatively late seasonal 
distribution is suggested, which may account for the apparent rarity of 
the species. 

Records. MEXICO: Sonora: Carbo, November, 10 (type series). 

Remarks. The type series of sonorae, collected in 1933 by H. S. Gentry, 
is in the collection of the California Academy of Sciences. In the holotype 
and one of the paratypes the elytral vittae begin at the humerus and 
extend to the apex; in the other specimens they are short of both the 
humerus and the apex. 

Lytta (Paralytta) biguttata LeConte 

Lijtta biguttata LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 332; 1866, Proc. Acad. Nat. Sci. Philadelphia, p. 349. Ulke, 1875, in Report 
upon the geographical and geological explorations and surveys west of the 
100th meridian, vol. 5, p. 825. Carruth, 1931, Ent. News, vol. 42, p. 54 (as 
biguttata Say). Tanner, 1934, Ann. Ent. Soc. America, vol. 27, p. 45. Vaurie, 
1951, Amer. Mus. Novitates, no. 1477, p. 43, fig. 14. Dillon, 1952, Amer. 
Midland Nat., vol. 48, p. 350. Selander, 1956, Bull. Brooklyn Ent. Soc, vol. 
51, p. 116. 

Cantharis biguttata, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 109. 
Snow, 1883, Trans. Kansas Acad. Sci., vol. 8, p. 43. Champion, 1892-93, Biol. 
Centrali-Americana, Coleoptera, vol. 4, pt. 2, pp. 445, 464, pi. 21, fig. 23. 
Wickham, 1896, Bull. Lab. Nat. Hist. State Univ. Iowa, vol. 3, p. 169 (as 
biguttata Say). Cockerell, 1898, New Mexico Agr. Exp. Sta. Bull. no. 28, 
p. 173; 1901, Ent. News, vol. 12, p. 40. Fall, 1901, Trans. Amer. Ent. Soc, 
vol. 27, p. 297. Wickham, 1902, Bull. Lab. Nat. Hist. State Univ. Iowa, 
vol. 5, p. 301. Fall and Cockerell, 1907, Trans. Amer. Ent. Soc, vol. 33, p. 210. 

Cantharis bivirgata Duges, 1881, La Naturaleza, vol. 5, p. 140, pi. 4, fig. 2. 



SYSTEMATICS AND PHYLOGENY 87 

Cantharis alfredi Duges, 1889, An. Mus. Michoacano, vol. 2, p. 98. New 
name proposed in error for bivirgata. 

Cantharis alemani Duges, 1889, An. Mus. Michoacano, vol. 2, p. 106. 

Cantharis bigitttata var. bivirgata, Champion, 1893, Biol. Centrali-Americana, 
Coleoptera, vol. 4, pt. 2, p. 464, pi. 21, fig. 24. 

Lytta biguttata pollens Selancler, 1956, Bull. Brooklyn Ent. Soc, vol. 51, 
p. 119, figs. 1-3, 4A. 

Lytta biguttata biguttata, Selancler, 1956, Bull. Brooklyn Ent. Soc, vol. 51, 
p. 112, figs. 1-3, 4B-D. 

Lytta biguttata bivirgata, Selancler, 1956, Bull. Brooklyn Ent. Soc, vol. 51, 
p. 125, figs. 2, 4E. 

Head varying from black (with an orange frontal spot) to completely 
orange. Antennae black, with basal segments sometimes orange. Prono- 
tum (Fig. 173) orange, finely margined with black; disk commonly with 
a black spot on each side just behind middle, sometimes also with a black 
dot on lateral margin before middle; discal spots sometimes extended to 
base of pronotum or fused, or both. Elytra (Fig. 160) yellow or orange, 
each with a brown or black spot or short vitta at apical fourth, or with a 
pair of black submarginal vittae, or immaculate; elytral suture usually 
finely margined with black for basal third to half (black margin occa- 
sionally broadened, fusing with discal spot). Under surface varying from 
largely black to entirely orange. Femora and tibiae varying from black 
to orange; tarsi black or brown. Wings varying from dark brown to nearly 
colorless. Pubescence silver or pale golden, conspicuous throughout. 
Length: 7 mm. -13 mm. 

Head smooth, shiny, sparsely to densely punctate; pubescence rather 
long, erect. Pronotum subcircular; basal margin more strongly recurved 
and much more deeply emarginate than in related species; disk strongly 
declivous basally; calluses poorly defined; surface smooth, shiny, more 
shallowly punctate than head, very sparsely punctate at center of disk; 
pubescence as on vertex. Elytra more weakly rugose than in related 
species; pubescence erect, variable in length. 

Male. Head behind eyes slightly flattened. Antennae (Fig. 76) reach- 
ing base of pronotum or nearly so; segment III weakly incrassate; IV-VI 
strongly incrassate; IV largest, clearly wider than distal segments, not 
curved; VII slightly incrassate, intermediate between VI and VIII in size 
and shape; VIII-X moniliform, longer than wide. Fifth abdominal ster- 
num shallowly emarginate. Sixth sternum as in scitula (see Fig. 234) or 
with emargination semicircular in form. Pygidium (Fig. 208) produced, 
oval, not at all acute. Genitalia as in Figure 306. 

Female. Sixth abdominal sternum truncate or shallowly, triangularly 
emarginate at apex (Fig. 279). 

Type locality. Of biguttata, Santa Fe, New Mexico; of alemani, [Ciu- 



88 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lljttd 

dad] Lerdo, Durango; of b. pallens, Douglas, Arizona; of bivirgata, Silao, 
Guanajuato. 

Geographic distribution. State of Mexico north to southern Utah and 
western Texas; thence north on the Great Plains to South Dakota and 
Montana. 

Seasonal distribution. March — October. 

Records. I have departed from my usual procedure in the present case 
by listing the records for the species by race. 

Typical race: UNITED STATES: Colorado: Antonito (Wickham, 
1902); Canon City (Wickham, 1902); Colorado Springs, June, 5; Cortez, 
July-August, 5; Denver, June-July, 4; Fort Collins (Wickham, 1902); 
near Hoehne, August, 1; Rocky Ford, September, 1; Sterling, July, 1; 
Trinidad, June-August, 4. Kansas: Cheyenne County, 3300 ft., July, 3; 
Dresden, July, 4; Gove County, 2813 ft., 3; Logan County, 3322 ft., 2; 
Quinter, July, 2; Wallace County, 3400 ft., July, 2. Montana: Phillips 
County, June, 1. Nebraska: Dix, July, 73; Mitchell, July, 1; Palisade, July, 
1. New Mexico: Bandelier, August, 1 (not located); Broadview, August, 
1; Cimarron, August, 1; Gallinas Canyon (Snow, 1883); Glorieta (Fall 
and Cockerell, 1907); near Hot Springs, Las Vegas, 7000 ft., August, 1; 
Koehler, 2; near Koehler, prairie, 1; Las Vegas, July, 6; Loving, May, 
September, 32; Maxwell, July, 6; Pecos ( Fall and Cockerell, 1907 ) ; Raton, 
August, 7; San Jose, July, 1; Santa Fe, July- August, 11; Torrance County, 
1. South Dakota: Ardmore, July, 5; Belle Fourche, July, 1; Cottonwood, 
June, 4; Highmore, June, 2; Haakon County (Carruth, 1931). Texas: 
Alpine, June-August, 4; The Basin, 5000 ft., Big Bend National Park, 
June, 1; Brady, May, 1; Chisos Mountains, Big Bend National Park, July, 
1; Cotulla, March-May, 44; Dalhart, June, 1; Davis Mountains, July, 1; 
100 mi. east of El Paso, July, 1; Fort Davis (Dillon, 1952); 20 mi. north 
of Fort Davis, 4000 ft., September, 1; 5 mi. north of Langtry, June, 1; 
Marfa, July, 17; Mason, May, 1; Odessa, May, 2; Pine Springs, Culbertson 
County, August, 1; Presidio, June-August, October, 27; Sanderson, May, 
2; 11 mi. north of Valentine, August, 5. Wyoming: 40 mi. north of Lusk, 
July, 1; Wheatland, August, 1. 

MEXICO: Coahuila: Cabos (Vaurie, 1950); Guadalupe, August, 1; 
La Gloria, south of Monclova (Vaurie, 1950); La Rosa (Vaurie, 1950); 
Paila (Vaurie, 1950); 5 mi. north of Saltillo, 5100 ft., August, 4; San 
Pedro [de Colonias], June, 1; 25 mi. southeast of San Pedro [de Colonias] 
(Vaurie, 1950); Torreon, June-August, 5. Durango: [Ciudad] Lerdo, 
May, August, 3; San Lucas (Vaurie, 1950); Yerbanis, Cuencame District 
(Vaurie, 1950). San Luis Potosi: El Huizache, 4500 ft., August, 1. 

Pale race: UNITED STATES: Arizona: 12 mi. west of Bisbee, 3; 
Camp Apache (Ulke, 1875); Carrizo Creek, highway 60, Navajo County, 



SYSTEMATICS AND PHYLOGENY 89 

August, 1; Cave Creek, Chiricahua Mountains, 5000 ft.-6000 ft., August, 1; 
Chiricahua Mountains, August, 4; Copper Basin, near Prescott, 1; Corn- 
ville, September, 1; near Del Rio Verde River, Yavapai County, Septem- 
ber, 2; Dewey, 1; Douglas, July, September-October, 18; Douglas to Chi- 
ricahua Mountains, September, 2; near Fort Whipple (LeConte, 1866); 
Gila Valley, Graham County, 1; Hereford, August, 5; 10 mi. southwest of 
Kayenta, 6000 ft., August, 4; Lochiel Station, August, 3; Mormon Lake, 
6900 ft., August, 1; Nogales, September-October, 4; Onion Flat, Chirica- 
hua Mountains, August, 1; Peach Springs, Hualpai Reservation, Septem- 
ber, 2; 7 mi. east of Pearce, August, 2; Pinery Canyon, 5000 ft., west slope 
Chiricahua Mountains, August, 1; Prescott, August, 8; San Bernardino 
Ranch, 3750 ft., Cochise County, August, 9; Santa Rita Mountains, July, 
1; Seligman, 4; Senator Mine, near Prescott, September, 1; Sierra Ancha 
Mountains, Gila County, July, 9; Springerville, August, 1; Yarnell, Sep- 
tember, 4; Walnut [National Monument], 1; Washington Mountains, near 
Nogales, September, 1; White Mountains, September, 1; Williams, August, 
1. New Mexico: 21 mi. southeast of Alamagordo, August, 3; Albuquerque, 
August, 3; Artesia, July, 12; Cambray, September, 9; Carlsbad, August, 
2; Deming, August, 3; 33 mi. east of Deming, 4300 ft., August, 2; Fort 
Wingate, October, 1; 18 mi. east of Gallup, August, 8; Las Cruces, June, 
September, 3; 49 mi. west of Las Cruces, August, 2; 10 mi. east of Las 
Cruces, 4400 ft., September, 1; Mescalero Reservation, September, 2; 
Mesilla Valley (Cockerell, 1898); Mule Creek, Grant County, 2; Roswell 
(Fall and Cockerell, 1907); Silver City, August, 5; Socorro (Cockerell, 
1898); White Sands Proving Ground, 4000 ft., September, 2. Texas: El 
Paso, 1; McNary, May, September, 5. Utah: Pintura, August, 3; St. 
George, 2; Zion National Park, 10. 

MEXICO: Chihuahua: 20 mi. east of San Miguel, Sonora, August, 3; 
20 mi. northwest of Nueva Casas Grandes, August, 2. 

Southern Mexico race: MEXICO: Guanajuato: State label only, 1; 
Silao (Duges, 1881); Tupataro (Duges, 1889); 1 mi. east of Villagran, 
July, 2. Mexico: 14 mi. west of Toluca, July, 1. 

Remarks. The present treatment is abbreviated as I have already 
presented an analysis of variation in this species, together with syno- 
nymical notes and distribution maps, in another paper (Selander, 1956). 
In the paper cited, three subspecies of biguttata were recognized. Such 
an arrangement seems to be useful, at least for the present time, but I 
now prefer to use vernacular, rather than scientific, names for the races. 
I therefore propose to substitute the name "typical race" for b. biguttata, 
"pale race" for b. pollens, and "southern Mexico race" for b. bivirgata. 

At the time my paper was published there was a wide distributional 
gap between the typical race in Coahuila and Durango and the southern 
Mexico race in the states of Guanajuato and Mexico. Since then I have 



90 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytttt 

received a female of biguttata collected August 22, 1954 by J. G. Chill- 
cott at El Huizache, in central eastern San Luis Potosi. This specimen 
differs from all others that I have seen in that the lateral lobes of the 
sixth abdominal sternum are acutely produced and the emargination of 
the base of the pronotum forms a deep, slotlike incision. It has the anten- 
nae and elytral pubescence of the typical race; on this basis I have 
assigned it to that race. In color it differs from the typical race in having 
the black discal mark on each elytron much broadened and extending 
from before the middle to near the apex, as in some specimens of the 
southern Mexico race. It is otherwise heavily marked in that the discal 
spots of the pronotum extend to the base and the legs are entirely black. 

The type of biguttata is in the Museum of Comparative Zoology. The 
type of b. pollens is in the collection of the University of California at 
Berkeley (examined). The type of bivirgata having been lost, I have 
designated as neotype a specimen labeled "Guanajuato, F. Duges, Samml. 
CI. Miiller," in the Zoologische Staatssammlung in Munich. This speci- 
men belongs to the typical form as described by Duges. The type of 
alemani cannot be located, but pending the opportunity of studying 
topotypical material, I have refrained from designating a neotype. 

Bionomics. This species occurs on a variety of Compositae. According 
to label data, specimens of the typical race have been collected on 
Applopappus spinulosus, Bahia absinthifolia, B. pedata, and Bailey a 
multiradiata at Presidio, Texas; on guayule (flowers) at Loving, New 
Mexico; on "sunflowers" at Quinter, Kansas; and on Verbesina encelioides 
at Cotulla, Texas. I have collected specimens on Hymenoxys richardsoni 
near Gallup, New Mexico, and on an unidentified composite near Villa- 
gran, Guanajuato. 

In addition to the above, there is a record of a single specimen from 
Petalostemum ( Leguminosae ) at San Jose, New Mexico, and Cockerell 
(1901) recorded specimens from Cleome serrulata ( Capparidaceae ) at 
Las Vegas, New Mexico. Both records should be regarded as question- 
able unless verified. 

Duges ( 1881 ) recorded a host plant for the southern Mexico race un- 
der the vernacular name "sin collar." This plant's identity is unknown 
to me. 

Lytta (Paralytta) scitula (Champion) 

Cantlxaris scitula Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 446, pi. 20, fig. 25. 

Cantharis signaticollis Champion, 1892, Biol. Centrali-Americana, Cole- 
optera, vol. 4, pt. 2, p. 446, pi. 20, fig. 24. New synonymy. 

Head black with an orange frontal spot ( rarely with labrum and under 
side of head partially orange also). Pronotum varying from orange with 



SYSTEMATICS AND PHYLOGENY 91 

a fine black margin and a pair of large black discal spots near middle 
to entirely black. Scutellum black. Elytra black with lateral margin 
and a broad discal vitta from base to near apex varying from yellow ( Fig. 
161 ) to a dark pitchy brown which is just perceptibly lighter in shade 
than rest of elytra. Wings light brown. Legs varying from orange with 
apex of femora and distal tarsal segments infuscate to entirely black. 
Under surface of thorax black. Abdomen varying from orange with first 
four sterna black except at sides to entirely black. Pubescence pale 
golden. Length: 7 mm. -10 mm. 

Head with vertex smooth, shiny, moderately coarsely, deeply, moder- 
ately densely punctate; surface of frontal area similar but punctures 
becoming denser, rather crowded; pubescence moderately long, erect. 
Pronotum subcircular, as long as wide; basal margin distinctly emarginate 
on midline, but not so deeply as in biguttata; midline impunctate before 
middle; discal calluses large, weakly elevated; surface and pubescence as 
on vertex. Elytral surface finely rugose reticulate; pubescence moderately 
long, semierect, conspicuous with magnification, denser on apical half 
than on basal half. 

Male. Head behind and above eyes deeply impressed; impression 
following dorsal margin of eye forward to the extent that side margins 
of head above eyes are distinctly concave in frontal view. Antennae 
(Fig. 77) reaching base of pronotum; segment III moderately incrassate, 

IV and V strongly so, VI weakly so, IV and V distinctly wider than 
following segments; IV largest, concave on inner side, weakly curved; 

V flat on inner side, not curved; VI slightly wider than VII; VII-X 
longer than wide, progressively slightly wider. Fifth abdominal sternum 
deeply, broadly emarginate. Emargination of sixth sternum (Fig. 234) 
very deep, broadly triangular; sternum impressed at center; lateral lobes 
of sternum acutely produced. Pygidium produced as in Figure 208. Geni- 
talia as in Figure 307. 

Female. Antennae as in quadrimaculata, not attaining base of prono- 
tum. Sixth abdominal sternum ( Fig. 280 ) truncate at apex, more strongly 
cupped than in any related species except quadrimaculata and proteus. 

Type locality. Of scitula, Chilpancingo [de los Bravos], 4600 ft., Guer- 
rero; of signaticollis, Matamoros Izucar [Izucar de Matamoros], Puebla. 

Geographic distribution. Southwestern Puebla, eastern Guerrero, and 
northwestern Oaxaca. (See Fig. 23.) 

Seasonal distribution. Collected in May and September. 

Records. MEXICO: Guerrero: Chilpancingo [de los Bravos], 4600 ft. 
(Champion, 1892). Oaxaca: Nochixtlan, September, 2. Puebla: Amozoc, 
July, 9; Atlixco, May, 6; Matamoros Izucar [Izucar de Matamoros], 2 
(paratypes of signaticollis); [Izucar de] Matamoros, September, 11. 



92 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 




Fig. 23. Distribution of Lytta scitula (circles) and Lytta sonorae (square). 



Remarks. The presence of a deep concavity on each side of the head 
in the male distinguishes scitula from all other species of Lytta except 
scituloides. In the latter species the concavity, although actually deeper 
than in scitula, does not extend far forward enough to affect the outline 
of the head in frontal view. In other respects scitula and scituloides are 
not particularly similar, as is indicated by the fact that they have been 
assigned to different subgroups in the present classification. 

It is of interest to note that the modification of the head of males 
of scitula and scituloides is paralleled in males of the Old World genus 
Oenas Latrielle. In Oenas, however, the concavity is much deeper and is 
rather sharply margined. 

The darkest and the palest combinations possible within the range of 
color variation described above are represented among specimens studied. 
Between these extremes a number of intermediate combinations are repre- 
sented, indicating that the variation is continuous. 

Scitula was originally described from Guerrero on the basis of a male 
having the pronotum and abdomen black, the elytra yellow vittate, and 
the legs partially orange. This specimen is in the British Museum (Natural 
History). Drawings of its genitalia have been made available to me 
through the courtesy of C. M. F. von Hayek. 



SYSTEMATICS AND PHYLOGENY 93 

Champion's description of signaticollis was based on four specimens 
from Puebla having the pronotum partially orange, the elytra entirely 
black or with the lateral margin yellow, and the legs and abdomen black. 
The type, a male, is also in the British Museum (Natural History). 
Included among the material I have studied are two paratypes of sig- 
naticollis, including the male figured in the Biologia Centrali-Americana. 

Bionomics: The series from Amozoc, Puebla, was collected on flowers 
of a species of Compositae which has not as yet been identified. The 
beetles were feeding on pollen. 

Lytta (Paralytta) qiiadrimaculata (Chevrolat) 

Cantharis qiiadrimaculata Chevrolat, 1834, Coleopteres du Mexique, fasc. 
4, p. [17]. Duges, 1869, La Naturaleza, vol. 1, p. 108; 1889, An. Mus. 
Michoacano, vol. 2, p. 99. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 105. 
Champion, 1892, Biol. Centrali-Americana, Coleoptera, vol. 4, pt. 2, p. 440, 
pi. 20, fig. 14. Fall, 1901, Trans. Amer. Ent. Soc., vol. 27, p. 296. 

Cantharis octomaculata, Pefiafiel y Barranco, 1866, Gaceta Med. Mexico, 
vol. 2, p. 225, pi. 1. Jimenez, 1866, Gaceta Med. Mexico, vol. 2, p. 228. 

Cantharis fasciolata Jimenez, 1866, Gaceta Med. Mexico, vol. 2, p. 253, 
pi. Duges, 1869, La Naturaleza, vol. 1, p. 107, pi. 1, fig. 5. 

Cantharis hifasciatus Duges, 1869, La Naturaleza, vol. 1, p. 106. 

Cantharis qiiadrimaculata var. bifasciata, Champion, 1892, Biol. Centrali- 
Americana, Coleoptera, vol. 4, pt. 2, p. 441, pi. 20, figs. 15-16. 

Lytta fasciolata, Denier, 1935, Rev. Argentina Ent., vol. 1, p. 25. 

Lytta qiiadrimaculata, Vaurie, 1950, Amer. Mus. Novitates, no. 1477, p. 41, 
fig. 13. Dillon, 1952, Amer. Midland Nat., vol. 48, p. 349. Selander, 1959, 
Proc. Ent. Soc. Washington, vol. 61, p. 208. 

Head black, with sides of vertex sometimes in part orange in the male. 
Pronotum yellowish orange, finely margined with black, with a large black 
discal spot on each side near middle and almost always a small black dot 
on lateral margin of disk just before middle; discal spots sometimes ex- 
tending to base of pronotum as thick vittae. Elytra usually yellowish 
orange with four black or brown spots of variable size and shape (Fig. 
159), rarely entirely yellowish orange or entirely black. Under surface of 
thorax black. Legs varying from entirely black to orange with apex of 
femora black. Abdomen entirely black or with some or all of the terga 
and sides of sterna orange; orange areas of sterna progressively larger 
toward apex of abdomen, those of fifth sternum sometimes meeting 
broadly on midline. Wings brown. Pubescence varying from silver 
throughout to black throughout. Length: 10 mm. -21 mm. 

Head distinctly wider than long; tempora rather prominent; surface 
smooth, shiny, moderately coarsely, deeply, moderately densely punctate; 
pubescence erect. Pronotum subcircular to nearly campanuliform, de- 
pending on degree of curvature of sides, about as long as wide; basal 
margin usually distinctly emarginate on midline; disk often broadly im- 



94 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lt/ttd 

pressed along midline, calluses weakly elevated; surface smooth, shiny, 
more finely, weakly, and almost always more sparsely punctate than ver- 
tex; center of disk usually subimpunctate; pubescence shorter and less 
conspicuous than on vertex. Elytra finely rugose reticulate; base rela- 
tively smooth, very finely punctate; posteriorly the punctures are gradu- 
ally enlarged to form cells between weak reticulations; pubescence short, 
very sparse, inconspicuous. 

Male. Head behind eyes flattened or very weakly concave. Antennae 
(Fig. 78) extending one or two segments beyond base of pronotum; seg- 
ments III-VII strongly incrassate; IV largest; V-VII progressively nar- 
rower; VIII-X much narrower, longer than wide. Fifth abdominal 
sternum shallowly emarginate. Emargination of sixth sternum (Fig. 235) 
very deep, broadly triangular; sternum flattened at center; lateral lobes 
of sternum acutely produced. Pygidium strongly and narrowly prolonged 
at middle, the projection variable in shape (Fig. 209). Genitalia as in 
Figure 308; ventral hooks of aedeagus unusually variable. 

Female. Sixth abdominal sternum truncate at apex; margin turned 
strongly dorsad so that sternum is strongly cupped (as in scitula and 
proteus). Pygidium rounded. 

Type locality. Of quadrimaculata, Orixaba [Orizaba, Veracruz]; of 
octomacidata, either Mineral del Chico or Atotonilco el Grande, both in 
Hidalgo; of fasciolata and bifasciatus, Mexico. 

Geographic distribution. Widespread at elevations between 4500 ft. 
and 9000 ft. on the Central Plateau of Mexico and adjacent southern 
highlands from Jalisco, Michoacan, Mexico (state), Puebla, and central 
western Veracruz north to southern Chihuahua and the Rio Grande. 
( See Fig. 24. ) 

Records of quadrimaculata from Texas are based on specimens col- 
lected many years ago. Horn (1873) mentioned specimens "said to have 
been collected in Texas near the Mexican border." Dillon's ( 1952 ) record 
is based on an old specimen in the United States National Museum 
labeled "Tex 6766, Collection C. V. Riley." In the absence of more recent 
records there is some doubt in my mind as to whether quadrimaculata 
actually occurs north of the Rio Grande, but inasmuch as there are reli- 
able records from Nuevo Leon and Coahuila, there is no real reason to 
ignore the Texas records. 

Vaurie (1950) erred in the identification of two of the localities re- 
corded for quadrimaculata by earlier workers. Thus "pueblo del Puente" 
(given as "El Puente" by Vaurie) of Duges (1869) is a locality in Hi- 
dalgo, not Nayarit, and the Almolonga of Champion ( 1892 ) is in all 
probability the settlement of that name in Veracruz, not that in southern 
Oaxaca. 



SYSTEMATICS AND PHYLOGENY 



95 



Seasonal distribution. May 23 — October 29. 

Records. UNITED STATES: Texas: State label only, 1; "near the 
Mexican boundary" (Horn, 1873). 

MEXICO: Country label only, 20. Aguascalientes: El Retono, 10 mi. 
east of Aguascalientes, August, 1; 4 mi. southwest of Aguascalientes, 
6100 ft., October, 1. Chihuahua: Salaices, 5200 ft, September, 1. Coa- 
huila: 15 mi. south of Arteaga, July, 1. Distrito Federal: Chapultepec, 
Coapa, Guadalupe, San Angel, Santa Fe, Tlalpam [Tlalpan], and district 
label only, July, August, October, 28. Durango: Durango, 10 (9 in July); 
6 mi. south of Durango, August, 1; San Lucas [Ocampo], 6700 ft., August, 
1; Sierra de Durango, 2; Villa [Ciudad] Lerdo, 8. Guanajuato: Guana- 
juato (Duges, 1889). Hidalgo: Atotonilco el Grande (Penafiel y Barranco, 
1866); Mineral del Chico (Penafiel y Barranco, 1866); Pachuca, May, 
June, 13; 16 mi. east of Pachuca, 7900 ft., June, 4; "pueblo del Puente" 
(Duges, 1869). Jalisco: Guadalajara, September, 7; La Punta, August, 4; 
4 mi. west of Mazamitla, 6800 ft., October, 11; Ojuelos de Jalisco, 7400 ft., 
August, 3; Sayula (Champion, 1892); Zapopan, September, 3; Zapotlan 
[Ciudad Guzman], 2. Mexico: Amecameca, June, 1; Atlacomulco, 8500 
ft., August, 2; Chapingo, September, 1; Cortez Pass, west slope, 9000 ft., 
July, 2; 4 mi. north and 10 mi. northwest of Ixtlahuaca, July, 60; San 




Fig. 24. Distribution of Lijtta quadrimaculata. 



96 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lyttd 

Rafael, 8100 ft., September, 8; Temescaltepec, 3; Tlalnepantla, Septem- 
ber, 1; Toluca and surrounding region (4 mi. east, 1.8 mi. southeast, 26 
mi. south, 14 mi. west, and 14 mi. and 15 mi. northwest), July, August, 
303; Zinacantepec, July, 8. Michoacdn: Morelia, September, 33; Tacam- 
baro [de Codallos] (Champion, 1892); Tancitaro, 8600 ft., August, 3. 
Nuevo Leon: 15 mi. southwest of China, July, 1. Pnebla: 5 mi. south of 
Acatzingo, July, 8; Amozoc, July, 42; Atlixco, 2; Esperanza, 10; Matamoros 
Izucar [Izucar de Matamoros] (Champion, 1892); Tecamachalco, July, 
1; Tehuacan, 2; 5 mi. northeast of Teziutlan, 4700 ft., June, 9. Queretaro: 
10 mi. east of San Juan del Rio, 6500 ft., 7. San Luis Potosi: Hacienda de 
Bleados [Rledos?], 4; San Luis Potosi, 7. Tlaxcala: 4 mi. and 10 mi. east 
of Apizaco, July, 16; 2.5 mi. west of El Carmen, June, 12; 4 mi. northwest 
and 9 mi. east of Huamantla, June, July, 20. Veracruz: Almolonga, 1; 
Jalapa, 1; Limon, July, 21; Orizaba, 9; Perote, 8500 ft., July, 42. Zacatecas: 
Ojocaliente, September, 1. 

Remarks. This is one of the most variable species of the genus Lytta. 
Variable characters include the color of the pubescence, legs, male 
antennae, and abdomen; elytral color pattern; shape of the male pygid- 
ium; and form of the aedeagus. In general the variation is geographical, 
with a considerable degree of concordance, tending to set off the popu- 
lation of Veracruz, Puebla, and Tlaxcala from the rest of the species. 

Although I originally prepared a detailed analysis of variation in 
quadrimaculata for inclusion in the present revision, so much material 
from critical areas has accumulated in the past three years that it seems 
preferable to delay publication of this until all the data can be incor- 
porated in it. I have described the first instar larva elsewhere ( Selander, 
1959). 

The present location of Chevrolat's type is unknown to me. In the 
cases of octomaculata and fasciolata, all the type material has been 
destroyed. As neotype of octomaculata I have designated a male in the 
collection of the California Academy of Sciences labeled "Pachuca, 8000 
ft., Hidalgo, July 1937, K. Maehler collection"; and as neotype of fascio- 
lata, a male in the British Museum (Natural History) labeled "Mexico, 
Salle collection." A type was probably never designated for bifasciatus as 
Duges credited the species to Sturm. There are a few specimens in the 
Duges collection at the Institute de Biologia in Mexico City that may 
belong to the series on which Duges' description was based, but there is 
no way of establishing this with certainty. A neotype designation seems 
to be in order, then, and I have designated the neotype of octomaculata 
as the neotype of bifasciatus also. 

Bionomics. Adults of this species are found on species of Papaveraceae 
and Compositae. The principal food plant seems to be Argemone platy- 



SYSTEMATICS AND PHYLOGENY 97 

ceras ( Papaveraceae ) , the common prickly poppy of the Central Plateau. 
I have collected the beetles on this plant at a number of localities in the 
states of Mexico, Puebla, and Tlaxcala. Adults also feed on other Papa- 
veraceae, as shown by the fact that in 1957 I found an individual feeding 
on Eschscholtzia in Coahuila. In the same year I also found the species 
feeding on both Argemone and a species of Compositae at Amozoc, 
Puebla. Two other records of Compositae as food plants are in the litera- 
ture: Pefiafiel y Barranco (1866) reported it from Viguiera dentata, and 
Duges (1869) reported it from Bidens lencantha. In the same paper 
Duges also recorded the species from a "Scymus." Chevrolat (1834) 
stated that his specimens were found on a species of "marguerite blanche." 
I have been unable to establish the identity of these last two plants. 

According to my observations, pollen is the primary food of adults 
of quadrimaculata, although some petal tissue is also eaten, at least on 
Argemone. The adult beetles are relatively sluggish: when disturbed 
they will sometimes fly, but their usual reaction is to crawl to the edge 
of the flower upon which they have been feeding and to drop to the 
foliage beneath or to the ground. Their color pattern offers fairly effective 
camouflage when they are feeding at the centers of Argemone flowers. 

Two other species of the Variabilis Group feed on Argemone; where 
the range of quadrimaculata overlaps the ranges of these, it is common 
to find individuals of quadrimaculata feeding on the same flowers with 
individuals of one or the other of these species. In the states of Puebla 
and Tlaxcala the species associated with quadrimaculata in this way is 
ebenina. In the state of Mexico it is plumbea. Both associates appear to 
have the same feeding habits and behavior as quadrimaculata except 
that they are slightly more active and alert. No antagonism between in- 
dividuals of different species has been observed, nor do the individuals 
of one species seem to be in the least disturbed by the presence of the 
others. In one instance I saw a male quadrimaculata mount a female 
ebenina and unsuccessfully attempt copulation. Normally, however, males 
confine their mating attempts to females of their species. 

In 1957 I obtained eggs from adults of quadrimaculata collected live 
July 14 at Amozoc, Puebla. The eggs were deposited sometime between 
July 20 and 24 (the exact date was not noted); they produced larvae 
July 30. 

Lytta (Paralytta) proteus Haag-Rutenberg 

Lytta proteus Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 24, p. 37. 

Cantharis sobrina Duges, 1881, La Naturaleza, vol. 5, p. 143, pi. 4, fig. 6; 
1889, An. Mus. Michoacano, vol. 2, p. 97. 

Cantharis bipartita Duges, 1889, An. Mus. Michoacano, vol. 2, p. 106. 

Cantharis proteus, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 448, pi. 20, figs. 22-23. 



98 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Black. Head with vertex and center of frontal area brownish orange 
varying to completely black except for an elongate orange frontal spot. 
Pronotum varying from brownish orange with a black discal spot on 
each side at middle to completely black. Elytra brownish orange with 
apex to as much as apical two-thirds black, or black with only humeri 
orange, or completely black. Wings dark brown. Pubescence piceous on 
head; usually paler, varying to silver or gold, on pronotum (especially 
basal half); silver on elytra and under surface, producing a dusty effect 
macroscopically where surface is black. Length: 8 mm. -16 mm. 

Head smooth, shiny, moderately coarsely, deeply, densely punctate: 
punctures not crowded except on lower frontal area; pubescence moder- 
ately long, rather silky, more or less erect. Pronotum as in Figure 174, 
usually longer than wide (rarely as wide as long); basal margin very 
shallowly emarginate at center; calluses large, strongly elevated; surface 
smooth, shiny, coarsely, densely punctate; pubescence as on vertex, tend- 
ing to slant toward callus on each side. Elytra rugose, impunctate; pubes- 
cence semierect, conspicuous, longer on sutural half of each elytron than 
on lateral half; average distance between setae near lateral margin about 
equal to length of a single seta. 

Male. Head behind eyes not noticeably flatter than in female. Anten- 
nae (Fig. 79) reaching or slightly surpassing base of pronotum; segments 
III— VII very weakly incrassate, as wide as or barely wider than VIII-X, 
which are subcylindrical. Fifth abdominal sternum moderately deeply, 
triangularly emarginate. Emargination of sixth sternum (Fig. 236) ex- 
tremely deep, U-shaped, extending nearly three-fourths length of ster- 
num; sternum impressed at center; lateral lobes of sternum strongly 
produced as slender processes, bluntly rounded at apex. Pygidium (Fig. 
210) weakly produced, acute at apex. Genitalia as in Figure 309; gono- 
styli in lateral view extremely broad, truncate; aedeagus with a single 
ventral hook; dorsal hook unusually massive. 

Female. Antennae as in Figure 125, not reaching base of pronotum. 
Sixth abdominal sternum truncate at apex; margin turned strongly 
dorsad so that sternum is strongly cupped (as in quadrimaculata and 
scitula ) . 

Type locality. Of proteus, Mexico; of sobrina, Penjamo, Guanajuato; 
of bipartita, Chilpancingo, Guerrero. 

Geographic distribution. Southern Nayarit and central southern Guana- 
juato to southeastern Guerrero, northern Oaxaca, and southwestern Pue- 
bla. ( See Fig. 25. ) 

Seasonal distribution. September 3 — December 2. 

Records. MEXICO: Country label only, 4. Colima: Colima, 2; Tonila, 
5. Distrito Federal: Mexico City (Champion, 1892); Tlalpan, September, 



SYSTEMATICS AND PHYLOGENY 



99 



3. Guanajuato: Penjamo, 1. Guerrero: Acahuizotla, 3500 ft., October, 4; 
Chilpancingo, 4600 ft., October, 2; Garciduena (not located) (Duges, 
1889); Hacienda de la Imagen (Champion, 1892); Rincon, 2800 ft., Octo- 
ber, 3; Taxco, October, 2; Tierra Colorado, 2000 ft., October, 2; Venta de 
Peregrino, 2. Jalisco: Guadalajara, 2 (1 in September); Zapopan, Septem- 
ber, 1. Mexico: [San Juan] Teotihuacan, November, 4; Temescaltepec, 12. 
Michoacdn: 7 mi. south of Arteaga, 3100 ft., December, 5; Jungapeo, 
September, 1; Tacambaro, October, 1. Morelos: Alpuyeca, 1100 m., Octo- 
ber, 2; Tepoztlan, 1750 m., September, October, 4. Nayarit: 4 mi. south 
of Santa Isabel, October, 26; 4 mi. northwest of Tepic, October, 51. 
Oaxaca: Huajuapan [de Leon], 1957 m., September, 1. Puebla: [Izucar 
de] Matamoros, 7 (3 in September); Puebla, 1. 

Remarks. While possessing the diagnostic features of the Quadrimacu- 
lata Subgroup, proteus is not particularly close morphologically to any of 
the other included species. The presence of only a single ventral hook 
on the aedeagus at first suggested a possible relationship between proteus 
and bipuncticollis ( in the Koltzei Subgroup ) , but a study of other char- 
acters has made it apparent that the loss of one of the hooks must have 
taken place independently in these two species. 

Although color variation in proteus is more or less continuous, it is 




Fig. 25. Distribution of Lytta proteus. 



100 



BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF LyttCl 



useful to treat it in terms of varieties, as outlined in Table III. These 
varietal categories readily accommodate the vast majority of specimens 
examined; only occasionally has it been necessary to make an arbitrary 
assignment. 

As shown in Table III, specimens with a black pronotum (varieties 
a and b) and those with an orange pronotum (varieties c through /) are 
about equally represented in the material studied. Some correlation 
exists between the color of the pronotum and that of the elytra; this is 
indicated by the fact that a good majority of the specimens with a black 
pronotum have entirely black elytra, while a majority of those with an 
orange pronotum have some orange on the elytra as well. Among the 
specimens with an orange pronotum, the vertex varies in color regardless 
of the color of the elytra. Unexpectedly, the vertex is orange in a larger 
proportion of the specimens with entirely black elytra than of those with 
some orange on the elytra. It is noteworthy that the vertex is always 
black when the pronotum is black, with the result that two of the eight 
possible color combinations are eliminated. 

From the data contained in Table III it is concluded that the genetic 
basis of color variation in proteus is fairly complex and that an interaction 
of factors is probably involved. 

When analyzed on the basis of samples from individual states (Table 
IV), color variation of the head, pronotum, and elytra, whether treated 
separately or in terms of the various combinations represented by the 
varieties just discussed, seems to exhibit no significant correlation with 
geographic distribution. Many additional data are needed, however, 
before this point can be established with certainty. 

As lectotype of proteus I have designated a specimen of variety / in 
the Zoologische Staatssammlung in Munich ( Haag-Rutenberg collection ) . 



Table III. Color Varieties of Lytta proteus and Their Frequency Distribution 

in Material Studied 





COLOR 


NUMBER 


PERCENTAGE 




VARIETY 


OF SPECIMENS 


OF 


SPECIMENS 


Pronotum (and vertex) black 










Elytra black 


a 


56 




38.4 


Elytra in part orange 


b 


19 




13.0 


Pronotum orange, bimaculate 










Elytra black 










Vertex black 


c 


2 




1.4 


Vertex orange 


d 


16 




11.0 


Elytra in part orange 










Vertex black 


e 


29 




19.9 


Vertex orange 


f 


24 




16.4 



SYSTEM ATICS AND PHYLOGENY 101 

Table IV. Frequency Distribution of Color Varieties in Samples of Lytta proteus 



SAMPLE 


a 


b 


c 


d 


e 


/ 


Nayarit 


34 


6 






24 


13 


Jalisco 


2 




1 








Guanajuato 


1 












Colima 


4 


1 




1 




1 


Michoacdn 








1 




6 


Mexico and Distrito Federal 


5 


1 


1 


10 


2 




Morelos 


2 


4 










Puebla 


2 


5 






1 




Oaxaca 




1 










Guerrero 


4 


1 




4 


2 


2 


State unknown 


2 










2 



The types of sobrina and bipartita have been lost. As neotype of sobrina 
I have designated a male of variety a from Penjamo, Guanajuato. This 
specimen was formerly in the collection of Salle, to whom it was probably 
sent by Duges. As neotype of bipartita I have designated a male of variety 
/ from Rincon, Guerrero. Both neotypes are now in the British Museum 
(Natural History). 



RETICULATA GROUP 



Colors nonmetallic; elytra black. Head with or without a pale frontal 
spot. Head quadrate, about as long as wide; frontal area flattened; sur- 
face minutely granular, sparsely micropunctate. Male head behind eyes 
more densely micropunctate, more finely punctate, and duller than at 
center of vertex, clothed with shorter ( very short ) setae, but not flattened 
or impressed. Male antennae reaching base of pronotum or extending a 
little beyond; segments elongate; IV-VII weakly to strongly incrassate. 
Eyes broad, prominent. Pronotum subcircular or transversely oval; disk 
lacking calluses. Elytra strongly reticulate, minutely granular, dull, sub- 
impunctate, essentially glabrous. Legs unmodified. Tarsal pads moder- 
ately developed; pale pubescence (pad) of first segment of middle tarsi 
limited to apical half or third, of hind tarsi to extreme apex. Tarsal claws 
cleft to base, with blades subequal in length. Male fifth abdominal ster- 
num shallowly emarginate. Male sixth sternum shallowly sulcate on 
midline; emargination lined with short setae. Pygidium not produced. 
Male genitalia with gonostyli stocky, truncate or obtusely rounded at 
apex; gonocoxal plate with lateral margins extending mesad at base as 
short, blunt processes; aedeagus stocky, deep, with two ventral hooks of 
nearly equal size. 

Remarks. This group contains seven species, none of which is par- 
ticularly strongly differentiated morphologically from the others. These 



102 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LljttCl 

may be divided into two subgroups on the basis of characters of color 
and a rather subtle difference in the form of the gonostyli of the male 
genitalia. The distribution of the group is centered in the southwestern 
United States and adjacent northern Mexico, with an outlying species, 
corallifera, of more southern distribution in San Luis Potosi and Hidalgo 
(Fig. 4). 

The strongly reticulate, black elytra distinguish members of the 
Reticulata Group from all other members of the genus Lytta except three 
species (magister, vulnerata, and margarita) of the Magister Group. The 
relationships of the Reticulata Group are clearly with the Variabilis 
Group. As mentioned before, the two groups could easily be treated as 
a single unit, since they agree in most characters, including the peculiar 
modification of the head in the male. 

The only species of the Reticulata Group whose host plant relation- 
ships are known is reticulata, which feeds on Convolvulaceae. 

Key to Subgroups 

1. Pronotum unicolored; front of head between eyes entirely black or 
with at most a vaguely defined orange suffusion; male genitalia with 

gonostyli straight, not bowed ( as in Fig. 310a) 

Reticulata Subgroup ( p. 102 ) 

Pronotum bicolored; front of head between eyes largely orange or 
with a well-defined orange spot; male genitalia with gonostyli bowed 
(as in Fig. 314a) Deserticola Subgroup (p. 108) 

Reticulata Subgroup 

Front of head between eyes entirely black or with a vaguely defined 
orange suffusion (some arizonica). Pronotum unicolored. Legs entirely 
black. Male genitalia with gonostyli straight, not bowed. 

Remarks. This subgroup contains the species reticulata, agrestis, miri- 
fica, and arizonica. The last three of these appear to be quite localized 
in distribution and are rare in collections. 

Key to Species 

1. Entirely black; elytra coarsely reticulate reticulata (p. 103) 

Pronotum orange 2 

2. Head orange above eyes agrestis ( p. 105 ) 

Head entirely black 3 

3. Pronotum subcircular ( Fig. 175 ) ; elytra coarsely reticulate, most cells 
at least .5 mm. in diameter. Rio Grande region near El Paso, Texas . . . 
mirifica ( p. 107 ) 



SYSTEMATICA AND PHYLOGENY 103 

Pronotum transversely suboval (Fig. 176); elytra finely reticulate, 

most cells less than .5 mm. in diameter. Northwestern Arizona 

arizonica ( p. 108 ) 

Lytta (Paralytta) reticulata Say 

Lytta reticulata Say, 1824, Jour. Acad. Nat. Sci. Philadelphia, vol. 3, p. 305. 
Horn, 1872, Rept. U.S. Geol. Surv. 1871, p. 390. Dillon, 1952, Amer. Midland 
Nat, vol. 48, p. 351. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, 
p. 85. 

Cantharis reticulata, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. Ill; 
1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. 
Soc, vol. 27, p. 298. Milliken, 1921, U.S. Dept. Agr. Bull. no. 967, p. 5, fig. 2. 

Black or piceous. Head lacking a pale frontal spot. Wings dark brown. 
Length: 10.5 mm. -22.0 mm. 

Head alutaceous, moderately shiny, deeply, densely punctate; pubes- 
cence short, erect. Pronotum subcircular, barely to one-tenth wider than 
long; disk convex, often impressed on each side behind middle; surface 
sculpture and pubescence as on vertex except at center of disk, which is 
impunctate and glabrous, punctures denser behind middle than before. 
Scutellum densely punctate and pubescent. Elytra coarsely reticulate, 
most cells .5 mm. or more in diameter. Hind tibiae moderately bowed. 
Outer hind tibial spur no longer than inner spur, two to three times as 
wide. Pad of first segment of middle tarsi limited to apical third. 

Male. Antennae (Fig. 80) with intermediate segments only slightly 
incrassate; IV-VI no wider or longer than VII. Emargination of sixth 
abdominal sternum (Fig. 237) moderately deep, broadly triangular; 
lateral lobes of sternum well rounded. Genitalia as in Figure 310; gono- 
styli truncate. 

Female. Antennae as in Figure 127. 

Type locality. "Inhabits Arkansa" (Say, 1824). 

Geographic distribution. Great Bend section of Texas north through 
the Great Plains to western Nebraska and southwestern Wyoming. (See 
Fig. 26.) 

Inasmuch as there is no other evidence that this species ranges west 
of the Great Plains, I have disregarded two specimens in the Carnegie 
Museum ( Klages collection ) labeled Monticello, San Juan County, Utah, 
August 1920. 

Seasonal distribution. April (Dillon, 1952) — October. May 15 is the 
earliest collection date of any specimen examined. 

Records. UNITED STATES: Colorado: State label only, 2; Akron, 
July, 14; Fort Morgan, July, 2; 15 mi. west of Fort Morgan, July, 1; 
Fountain Valley School, Colorado Springs, August, 1; Hardin, August, 1; 



104 BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lytta 

Hudson, July, 1; La Junta, August, 1; Logan County, August, 1; Roggen, 
July, 1; Sterling, September, 1; Wray, July, 3. Kansas: State label only, 
7; Clark County, 1962 ft., June, 2; Garden City, June, July, 6; Gove 
County, 2500 ft., 2; Hamilton County, 3350 ft., August, 3; Seward County, 
July, 1; Syracuse, May, 2; Wallace County, 3000 ft., 3. Nebraska: 32 mi. 
south of Gordon, August, 5; "Sand Hills," September, 1. New Mexico: 
State label only, 1; 5 mi. east of Acme, August, 75; Portales, July, 12; 
Roosevelt County, July, 1; Tucumcari, June, 3. Oklahoma: State label 
only, 1; Woodward, June, 1. Texas: Amarillo, July, 1; Brownfield, May, 
4; Canadian, June, 1; Ector County, May, 5; Higgens, May, 1; Lamesa 
(Dillon, 1952); Presidio (Dillon, 1952). 

Remarks. This is the most widely distributed and most commonly 
collected species of the Reticulata Group. It is easily recognized since 
it is the only entirely black species of Lytta with reticulate elytra. 

The first instar larva of reticulata has been described by MacSwain 
(1956). Say's type has been lost. As neotype I have designated a male 
in the Snow collection, University of Kansas, from 32 mi. south of Gordon, 
Sheridan County, Nebraska. 



Fig. 26. Distribution of Lytta reticulata ( circles ) and Lytta deserticola ( squares ) . 



SYSTEMATICS AND PHYLOGENY 



105 



Bionomics. Milliken (1921) reported that in the course of his work 
at Garden City, Kansas, "adults were taken on the bush morning-glory 
(Ipomoea leptophijlla) , excepting one which was found on alfalfa." 
Included in the material from Garden City which I have studied is one 
of Milliken's specimens from bush morning-glory (Convolvulaceae), two 
specimens collected by him on "creeping bur," and a specimen without 
the collector's label from sugar beet. Dillon ( 1952 ) reported "wild morn- 
ing-glory" as a host plant in Texas. 

Lytta (Paralytta) agrestis (Fall) 

Cantharis agrestis Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 302. 
Lytta agrestis, Maehler, 1939, Pan-Pacific Ent., vol. 15, p. 65. 

Black. Vertex and upper frontal area of head, entire pronotum, and 
base of elytra reddish orange. Orange area of elytra not reaching lateral 
margins, extending farthest posteriad along subsutural nervule, generally 
conspicuous but sometimes almost piceous and rather poorly defined. 
Wings dark brown. Length: 13 mm.-17 mm. 




Fig. 27. Distribution of Lytta agrestis (circles), Lytta mirifica (square), and Lytta 
arizonica (triangle). 



106 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Head smooth, shiny, moderately coarsely, weakly, sparsely, irregularly 
punctate; pubescence short, erect, especially distinct against pale area. 
Pronotum subcircular, one-tenth or less wider than long; disk evenly, 
weakly convex, a shallow fovea on each side behind middle; surface 
sculpture and pubescence as on vertex. Scutellum having a few punc- 
tures and setae. Elytra moderately coarsely reticulate; reticulations inter- 
mediate in coarseness between those of reticulata or mirifica and deserti- 
cola or arizonica. Outer hind tibial spur longer than inner spur, three to 
four times as wide. Pad of first segment of middle tarsi limited to apical 
third or half. 

Male. Antennae (Fig. 81) with intermediate segments moderately in- 
crassate; V-X progressively shorter and narrower. Emargination of sixth 
abdominal sternum (Fig. 238) moderately deep, somewhat rounded; 
lateral lobes of sternum broadly rounded. Genitalia as in Figure 311; 
gonostyli truncate. 

Female. Antennae as in Figure 128. 

Type locality. Arizona. 

Geographic distribution. Painted Desert, in the Navajo country of 
northeastern Arizona. (See Fig. 27.) 

Seasonal distribution. July 13 — August 27. 

Records. UNITED STATES: Arizona: State label only, 6 (including 
2 cotypes ) ; Painted Desert, August, 2; Tuba City, 6 ( 1 in July ) ; 20 mi. 
east of Tuba City, July, 3. 

Remarks. The elytra give the impression of having been washed basally 
with some of the orange color of the pronotum. Heretofore, this feature 
has seemed diagnostic of agrestis, but it is now known to occur also, in a 
much less well-marked form, in some mirifica and arizonica. Moreover, 
variation in the shade and extent of the orange elytral marking in agrestis 
is such that individuals with entirely black elytra will probably be found 
eventually. 

The specimens from 20 mi. east of Tuba City, Arizona, are three of 
four males previously recorded by Maehler ( 1939 ) . These were collected 
July 26, 1937, and July 27, 1937, not June 26, 1938, as stated by Maehler. 
Two of the specimens are in the American Museum of Natural History; 
the other specimen is in the collection of the California Academy of 
Sciences. 

Agrestis was described by Fall from three specimens taken at an un- 
recorded locality in Arizona. Two of these specimens are in the United 
States National Museum. The type is in the Museum of Comparative 
Zoology. 



SYSTEMATICS AND PHYLOGENY 



107 



Lytta (Paralytta) mirifica Werner 
Lytta mirifica Werner, 1950, Psyche, vol. 57, p. 133, figs. 1, 3. 

Black. Head lacking a pale frontal spot. Pronotum dark reddish orange. 
Immediate base of elytra sometimes suffused with orange. Wings nearly 
colorless except for clouded, light brown apex. Length: 14 mm.-21 mm. 

Head alutaceous, moderately shiny, coarsely, deeply, sparsely to moder- 
ately densely punctate, subglabrous. Pronotum (Fig. 175) subcircular, 
not appearing transverse, averaging (eight specimens) .08 (.03-. 11) 
wider than long; sides rounded but with a suggestion of lateral angles 
just before middle; disk evenly, weakly convex, a shallow fovea or broad 
impression on each side behind middle; surface as on vertex but more 
regularly sculptured; punctures and setae confined mainly to sides. 
Scutellum rather narrow, rounded, moderately densely punctate, gener- 
ally foveate, clothed with short setae. Elytra coarsely reticulate, most 
cells .5 mm. or more in diameter. Outer hind tibial spur longer than 
inner spur, two to three times as wide. Pad of first segment of middle 
tarsi limited to apical third or half. 

Male. Antennae (Fig. 82) with intermediate segments slightly incras- 
sate; V-X progressively a little shorter and narrower. Emargination of 
sixth abdominal sternum ( Fig. 239 ) moderately deep, broadly triangular; 
lateral lobes of sternum broadly rounded. Genitalia as in Figure 312; 
some variation present in the shape of the gonocoxal piece; gonostyli ob- 
tusely rounded at apex. 

Female. Antennae as in Figure 129. Genitalia with each gonocoxite 
consisting of a single, undivided sclerite. 

Type locality. Anthony, New Mexico. 

Geographic distribution. Known only from the type locality, in the 
Rio Grande Valley. ( See Fig. 27. ) 

Seasonal distribution. May 23 — July 15. 

Records. UNITED STATES: New Mexico: Anthony, May-July, 90. 

Remarks. This species is very similar to its presumed nearest relative, 
arizonica. It differs mainly in having the pronotum more nearly circular 
in shape and of a slightly different color and the elytra more coarsely 
reticulate. Mirifica and arizonica are the only species of Lytta possessing 
the combination of black head, orange pronotum, and reticulate elytra. 

The only material of this species known at the present time is a series 
of more than 100 specimens collected at the type locality in 1941 by R. H. 
Crandall. The type is in the Museum of Comparative Zoology. 



108 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Lytta (Paralytto) arizonica Selander 
Lytta arizonica Selander, 1957, Proc. Ent. Soc. Washington, vol. 59, p. 141. 

Black. Head suffused on frontal area with dark orange or piceous. 
Pronotum yellow-orange. Immediate base of elytra sometimes suffused 
with orange. Wings uniformly dark brown. Length: 13 mm. -20 mm. 

Differs structurally from mirifica as follows. 

Pronotum ( Fig. 176 ) transversely oval, averaging ( 13 specimens ) .15 
(.11-.22) wider than long; sides more evenly rounded, particularly from 
middle to apex. Scutellum wider. Elytra rather finely reticulate, all or 
nearly all cells less than .5 mm. in diameter. 

Male. Genitalia with gonocoxal piece tending to be slightly shorter 
and more truncate. 

Female. Genitalia with anterior end of each gonocoxite separated from 
posterior portion by a membranous area. 

Type locality. Littlefield, Arizona. 

Geographic distribution. Valley of the Virgin River in northwestern 
Arizona. (See Fig. 27.) 

Seasonal distribution. The type series was collected April 20. 

Records. UNITED STATES: Arizona: Littlefield, April, 13 (type 
series ) . 

Remarks. The orange color at the base of the elytra, when present, is 
scarcely noticeable without close inspection. The same is true in mirifica. 
In agrestis, on the other hand, it is conspicuous even when least de- 
veloped. 

The type of arizonica is in the American Museum of Natural History 
( examined ) . 

Deserticola Subgroup 

Front of head between eyes largely orange or with a well-defined 
orange spot. Pronotum bicolored. Male genitalia with gonostyli bowed 
in dorsal or ventral view. 

Remarks. This subgroup contains the species cribrata, deserticola, and 
corallifera. The first of these agrees with the species of the Reticulata 
Subgroup in the form of the male sixth abdominal sternum and in having 
entirely black legs. Deserticola and corallifera differ from all other species 
of the Reticulata Group in having this sternum deeply, roundly emargi- 
nate and the femora largely orange. 

Key to Species 

1. Femora black; emargination of male sixth abdominal sternum moder- 
ately deep, broadly triangular; elytra finely reticulate; head black 



SYSTEMATICS AND PHYLOGENY 109 

with an orange frontal spot cribrata (p. 109) 

Femora orange with black apex and base; emargination of male sixth 

abdominal sternum deeper, rounded 2 

2. Elytra finely reticulate, most cells less than .5 mm. in diameter; head 
orange with black at sides of frontal area. Southwestern United States 

and Sonora deserticola (p. Ill ) 

Elytra coarsely reticulate, most cells at least .5 mm. in diameter; head 
with orange confined to dorsal margin (including tempora) and a 
large frontal spot. San Luis Potosi and Hidalgo. . .coralliferq (p. 112) 

Lytta ( Paralytta ) cribrata LeConte 

Lytta cribrata LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 447. Dillon, 1952, Amer. Midland Nat., vol. 48, p. 352. 

Cantharis cribrata, Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. Ill; 
1883, Trans. Amer. Ent. Soc., vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. 
Soc, vol. 27, p. 298. 

Cantharis intricata Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 441, pi. 20, fig. 18. New synonymy. 

Cantharis pilsbryi Skinner, 1906, Ent. News, vol. 17, p. 217. 

Lytta intricata, Valine, 1950, Amer. Mus. Novitates, no. 1477, p. 43. 

Black. Head with a diamond-shaped orange frontal spot which is 
three-fifths or more as wide as frontal area between eyes. Pronotum 
bordered at sides with orange, except at base and apex. Wings uniformly 
dark brown or very pale brown with darker costal region and apex. 
Length: 18 mm. -20 mm. 

Head smooth, shiny, very finely, weakly, very sparsely punctate, 
subglabrous; punctures slightly deeper and more distinct than in deserti- 
cola; tempora more quadrate. Pronotum subcircular, one-tenth to one- 
fifth wider than long; sides more evenly rounded than in deserticola; disk 
evenly convex, a small impression near each lateral margin at and behind 
middle; surface as on vertex but punctures confined to sides. Scutellum 
as in deserticola. Elytra rather finely reticulate, most cells less than .5 mm. 
in diameter. Outer hind tibial spur slightly longer than inner spur, about 
twice as wide. Pad of first segment of middle tarsi limited to apical half. 

Male. Antennae as in deserticola (see Fig. 83) but longer, reaching 
one or two segments beyond base of pronotum, and with segments IV-VI 
in some specimens not as strongly incrassate, no wider than distal seg- 
ments. Emargination of sixth abdominal sternum moderately deep, 
broadly triangular, as in mirifica (see Fig. 239). Genitalia as in Figure 
313; gonostyli truncate. 

Female. Antennae as in deserticola (see Fig. 130), but longer. 

Type locality. Of cribrata, vicinity of Eagle Pass, Texas; of intricata, 
Villa [Ciudad] Lerdo, Durango; of pilsbryi, High Bridge, Pecos River, 
Texas. 



110 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



Geographic distribution. Central Chihuahua and northeastern Durango 
northeast to the Chinati Mountains and Eagle Pass, Texas. ( See Fig. 28. ) 

Seasonal distribution. August 15 — October 14. Skinner's (1906) record 
of April 27 for the type series of pilsbryi seems doubtful, especially since 
there is no date given on the label of the cotype specimen I have studied. 

Records. UNITED STATES: Texas: Chinati Mountains, 6 (5 in Octo- 
ber ) ; High Bridge, Pecos River, 1 ( cotype of pilsbryi ) ; vicinity of Eagle 
Pass (LeConte, 1853). 

MEXICO: Chihuahua: 12 mi. northwest of Gran Morelos, August, 4; 
Ojinaga, October, 1. Durango: Villa [Ciudad] Lerdo, 2 (paratypes of 
intricata ) . 

Remarks. The black vertex and femora at once separate cribrata from 
both deserticola and corallifera. In addition, the pronotum is more ex- 
tensively marked with black than in either of these species. The form of 
the emargination of the sixth abdominal sternum of the male of cribrata 
is distinctive within the limits of the subgroup. 

Two paratypes of intricata from Durango differ from specimens of 
cribrata from Texas and Chihuahua only in that their wings are pale 
brown with darker costal region and apex rather than uniformly dark 




Fig. 28. Distribution of Lytta cribrata (circles) and Lytta corallifera (squares). 



SYSTEMATICS AND PHYLOGENY 111 

brown. According to C. M. F. von Hayek (in lift.), the wing color of 
the type of intricata is the same as that of the paratypes studied. 

On the basis of a cotype in the American Museum of Natural History, 
pilsbryi differs in no way from cribrata. It has already been reduced to 
synonymy by Dillon (1952). 

A specimen of Lytta said to approach the type of cribrata was recorded 
from 20 mi. east of Tuba City, Arizona, by Maehler (1939), but from 
distributional evidence it seems unlikely that this was the species actually 
represented. 

The type of cribrata is in the Museum of Comparative Zoology; the 
type of intricata is in the British Museum (Natural History); and the 
type of pilsbrifi is in the collection of the Philadelphia Academy of Sci- 
ences. 

Lytta (Paralytta) deserticola Horn 

Lytta deserticola Horn, 1870, Trans. Amer. Ent. Soc, vol. 3, p. 90. . 

Cantharis deserticola, Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. Ill; 
1883, Trans. Amer. Ent. Soc., vol. 10, p. 310. Wiekham, 1896, Bull. Lab. 
Nat. Hist. State Univ. Iowa, vol. 4, p. 311. Cockerell, 1898, New Mexico Agr. 
Exp. Sta. Bull. no. 28, p. 173. Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, 
p. 296. Fall and Cockerell, 1907, Trans. Amer. Ent. Soc, vol. 33, p. 210. 

Black. Vertex and frontal area of head orange; frontal area between 
eyes bordered at sides with black, each border at epistomal suture about 
one-third as wide as frontal area, becoming narrower above. Pronotum 
reddish orange, finely margined with black; two or four small black spots 
in a transverse row at middle of disk; spots variable in size, at one ex- 
treme reduced to faint dots, at the other fused to one another or with 
apical black border, or both; when four spots are present, mesal pair 
usually larger than lateral pair (Fig. 177). Wings dark brown. Femora 
reddish orange with broadly black base and apex. Basal black area of 
fore femora extending to, or nearly to, middle of segment; of middle 
femora somewhat smaller; of hind femora confined to extreme base of 
segment or absent. Length: 11 mm. -23 mm. 

Head smooth, shiny, very finely, weakly, very sparsely punctate, sub- 
glabrous. Pronotum (Fig. 177) subcircular, one-tenth or less wider than 
long; disk evenly convex, one or two shallow impressions on each side 
near middle; surface as on vertex but punctures confined to sides. Scutel- 
lum with a few fine punctures and setae and a median fovea or sulcus. 
Elytra rather finely reticulate, most cells less than .5 mm. in diameter. 
Outer hind tibial spur slightly longer than inner spur, about twice as 
wide. Pad of first segment of middle tarsi limited to apical half. 

Male. Antennae (Fig. 83) with intermediate segments moderately in- 
crassate; V-X progressively shorter and narrower. Emargination of sixth 



112 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

abdominal sternum (Fig. 240) deep, rounded; lateral lobes of sternum 
bluntly rounded, not convergent. Genitalia as in Figure 314; gonostyli 
truncate; gonocoxal piece unusually short. 

Female. Antennae (Fig. 130) with segments VI-VIII each barely 
shorter than IV or V; IX and X slightly shorter than VIII. 

Type locality. South of Fort Whipple, Arizona. 

Geographic distribution. Southern Sonora north through southeastern 
Arizona and southwestern New Mexico to southwestern Utah (Iron 
County). (See Fig. 26.) 

Seasonal distribution. July — September 28. 

Records. UNITED STATES: Arizona: Bisbee, 1; Chiricahua Moun- 
tains, August, 1; Cochise County, 1; south of Fort Whipple (Horn, 1870); 
Graham Mountains, 9000 ft., September, 1; Grand Canyon, September, 1; 
Grand Canyon Rim Camp Ground, August, 1; Grand Canyon, north rim, 
August, 1; Nogales, 2; 5 mi. west of Portal, 5400 ft., August, 1; San Ber- 
nardino Ranch, 3750 ft., Cochise County, August, 2; Santa Rita Mountains, 
1; Skull Valley, September, 1; Thatcher, September, 1. New Mexico: State 
label only, 10 (1 in July); Cooney, August, 2; Socorro County, 1; Walnut 
Creek, near Silver City (Cockerell, 1898). Utah: Antelope Springs, Iron 
County, 2. 

MEXICO: Sonora: State label only, 3; Rio Mayo, September, 1. 

Remarks. The orange vertex and femora and the more deeply, roundly 
emarginate sixth abdominal sternum of the male easily distinguish this 
species and corallifera from other members of the Reticulata Group. 
From corallifera, deserticola is separated primarily by its more finely 
reticulate elytra, by differences in the color pattern of the head and 
pronotum, and by the form of the emargination of the sixth abdominal 
sternum of the male. 

The type of deserticola is in the collection of the Academy of Natural 
Sciences of Philadelphia. 

Lytta (Paralytta) corallifera Haag-Rutenberg 

Lytta corallifera Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 24, 
p. 35. 

Cantharis corallifera, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 441, pi. 20, fig. 17. 

Black. Tempora and top of head (except at midline) reddish orange; 
frontal area with an elongate orange spot which is three-fifths as wide 
as frontal area between eyes and is narrowly produced almost to epistoma. 
Pronotum bordered at sides with reddish orange, as in cribrata, but 
borders extending onto base of pronotum, where they are separated only 
by a narrow black stripe on midline. Wings pale brown, with darker costal 



SYSTEM ATICS AND PHYLOGENY 113 

region and apex. Femora reddish orange with broadly black base and 
apex; basal black area larger and more variable in size than in deserti- 
cola; orange area of fore femora sometimes more extensive than that of 
middle femora. Hind tibiae sometimes in part orange. Length: 16 mm.- 
23 mm. 

Head smooth, shiny, very finely, weakly, very sparsely punctate, sub- 
glabrous; punctures more distinct than in deserticola. Pronotum sub- 
circular, one-tenth to nearly one-fifth wider than long; disk evenly 
convex, several small impressions near each lateral margin at and behind 
middle; surface as on vertex but punctures confined to sides. Scutellum 
as in deserticola. Elytra coarsely reticulate (as in reticidata and rniri- 
fica), most cells .5 mm. or more in diameter; basal sixth of elytra not or 
only feebly reticulate. Outer hind tibial spur slightly longer than inner 
spur, two to three times as wide. Pad of first segment of middle tarsi 
limited to apical half. 

Male. Antennae as in deserticola (see Fig. 83). Emargination of sixth 
abdominal sternum ( Fig. 241 ) very deep, definitely semicircular, at 
least as wide as long; lateral lobes of sternum narrow, convergent. Geni- 
talia as in Figure 315; gonostyli truncate. 

Female. Not studied. 

Type locality. Mexico. 

Geographic distribution. Central Plateau of Mexico from southwestern 
San Luis Potosi to northern Hidalgo. ( See Fig. 28. ) 

Seasonal distribution. The single specimen studied for which seasonal 
data were recorded was collected August 27. 

Records. MEXICO: Country label only, 2. Hidalgo: Jacala, August, 1. 
San Luis Potosi: Sierra de San Miguelito, 1. State unknown: "Alton, s., 
ex Coll. J. Sturm," 1. 

Remarks. The female of this species apparently has not been collected. 
The specimen from Sierra de San Miguelito is one of four studied by 
Champion. As lectotype I have designated a specimen in Zoologische 
Staatssammlung in Munich ( Haag-Rutenberg collection ) labeled "Mexico, 
Baden." 

MAGISTER GROUP 

Head when black always with a pale frontal spot. Head and pronotum 
subglabrous, those setae present very short, erect. Male head not modi- 
fied behind eyes. Male antennae reaching at least as far as base of 
pronotum; male intermediate segments (IV-VII or VIII) weakly to 
strongly incrassate; distal segments slightly longer than wide. Eyes nar- 
row. Pronotum hexagonal to subcircular, lacking calluses. Elytra either 
rugose or reticulate; surface impunctate, essentially glabrous. Male 



114 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

middle tibiae sometimes impressed apieally. First segment of male 
middle tarsi never modified. Male fifth abdominal sternum shallowly 
emarginate or entire. Pygidium often produced, especially in male (al- 
ways distinctly produced if pronotum is not hexagonal). Aedeagus with 
two ventral hooks, the second one sometimes greatly enlarged. 

Remarks. This group contains nine species, divided equally into three 
subgroups. Its range extends from the peninsula of Baja California and 
the extreme northwestern corner of mainland Mexico to British Columbia 
(Fig. 3). 

In the adult stage the Magister Group is most clearly distinguished 
from the Variabilis and Reticulata groups by the absence of modification 
of the male head behind the eyes. From the Fulvipennis Group it is 
distinct in having the pronotum hexagonal or the male pygidium pro- 
duced, or both. This same combination of characters will distinguish it 
from the species of the Nitidicollis Group except moesta, which is at 
once distinct in having the blades of the tarsal claws fused. The known 
larvae of the Magister Group are separable from all other Lyttini on the 
basis of characters of the maxillary palpi (MacSwain, 1956). 

Within the Magister Group MacSwain's larval studies have demon- 
strated the presence of a major subdivision between magister and vul- 
nerata on the one hand and morosa, funerea, and blaisdelli on the other. 
The first group corresponds to my Magister Subgroup, which also in- 
cludes margarita, the larva of which is unknown. Within the second 
group of species MacSwain has shown that the larvae of morosa and 
blaisdelli share more similarities with each other than either does with 
the larva of funerea. However, judged by details of adult morphology, 
morosa and funerea appear to be much more closely related to each 
other than either is to blaisdelli. In the present instance I think the evi- 
dence of adult morphology is the more conclusive, and it is suggested 
that the similarities of the larvae of morosa and blaisdelli not shared by 
funerea are, in the main at least, primitive characters once possessed 
also by funerea but modified in the course of its evolution from an imme- 
diate common ancestor with morosa. 

Key to Subgroups 

1. Pronotum strongly hexagonal, with prominent, sharp lateral angles 
at middle (Figs. 183-84); head and pronotum usually at least in part 
orange; elytra coarsely reticulate or rugose; inner side of tibiae 
glabrous, except for a fine line of coarse setae along one or both edges 

Magister Subgroup (p. 127) 

Pronotum not as above, the lateral angles obtuse when present; pro- 
notum black; elytra rugose; tibiae normally pubescent on inner side, 
lacking lines of setae along inner edges 2 



SYSTEMATICS AND PHYLOGENY 115 

2. Male middle tibiae each with a deep, impunctate, glabrous impres- 
sion on anterior side apically; pronotum campanuliform or subeircular, 
without lateral angles (Figs. 181-82), always smooth; pygidium dis- 
tinctly produced Melaena Subgroup (p. 122) 

Male middle tibiae unmodified; pronotum subhexagonal, with obtuse 
lateral angles (Figs. 178-80), usually granular and alutaceous; py- 
gidium not produced; male genitalic type distinctive ( see below ) . . . 
Tenebrosa Subgroup ( p. 115 ) 

Tenebrosa Subgroup 

Entirely black (except for pale frontal spot) or with vertex pale. 
Head and pronotum usually granular and alutaceous. Pronotum sub- 
hexagonal, as wide as or wider than long, with obtuse lateral angles (Figs. 
178-80). Scutellum rounded. Elytra rugose. Tibiae normally pubescent, 
lacking lines of setae along inner edges. Middle tibiae less strongly 
bowed than in other subgroups. Male middle tibiae not impressed apically. 
Hind tarsal pads represented only by some pale pubescence near edges of 
ventral side of second to fourth segments. Male fore and middle tarsal 
pads well developed, dense, both larger and denser than in female. 
Pygidium rounded, not distinctly produced. Male genitalia with apical 
foramen of median tube extending to or nearly to apex of aedeagus; 
dorsal sclerite of aedeagus relatively slender, not forming a heavy, 
strongly recurved hook. 

Remarks. In the adult stage this subgroup is the most distinctive of 
the three comprising the Magister Group. It contains the species tene- 
brosa, blaisdelli, and lugubris. The first of these is quite different from 
the other two both morphologically and geographically and could rea- 
sonably be placed in a separate subgroup. The range of the subgroup is 
centered in California, with an extension along the Sierra Nevada into 
Oregon and thence eastward to southwestern Idaho. The host plant 
families recorded for the subgroup are Compositae (tenebrosa) and 
Scrophulariaceae ( blaisdelli ) . 

Key to Species 

1. Pronotum nearly quadrate, widest near apical margin (Fig. 178); 
segments IV-VII of male antennae with erect pale setae on ventral 
side; male sixth abdominal sternum with lateral lobes not produced 
( Fig. 242); ventral hooks of aedeagus subequal ( Fig. 316c); hind tibial 

spurs distinctly dissimilar in form; head variable in color 

tenebrosa ( p. 116 ) 

Pronotum more nearly hexagonal, widest near- middle (Figs. 179-80); 
male antennae not modified as above; male sixth abdominal sternum 



116 BIONOMICS, SYSTEM ATICS, AND PHYLOGENY OF Lytta 

with lateral lobes produced, hemispherical (Fig. 243); second ventral 
hook of aedeagus greatly enlarged (Figs. 317c-18c); head black with 

a small orange frontal spot between eyes 2 

2. Male genitalia as in Figure 317; wings pale brown, with at least apex 
darker, more opaque; one or both spurs of each hind tibia flattened, 
sticklike (Fig. 195); female sixth abdominal sternum with hind margin 

gradually turned dorsad blaisdelli (p. 119) 

Male genitalia as in Figure 318; wings colorless; both spurs of each 
hind tibia obliquely truncate (Fig. 194); female sixth abdominal 
sternum with hind margin turned abruptly dorsad, at a right angle . . . 
lugubris ( p. 120 ) 

Lytta (Paralytta) tenebrosa (LeConte) 

Cantharis tenebrosa LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 160. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 108; 1874, Trans. 
Amer. Ent. Soc, vol. 5, p. 38. Fall, 1901, Occas. Papers California Acad. 
Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc, vol. 27, p. 297. 

Lytta tenebrosa, LeConte, 1853, Proc Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 333. 

Cantharis occipitalis Horn, 1883, Trans. Amer. Ent. Soc, vol. 10, p. 312. 
Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 298. New synonymy. 

Lytta occipitalis, Linsley and MacSwain, 1942, Amer. Midland Nat., vol. 
27, p. 406, tbl. 1, fig. 6. 

Black. Head with an orange frontal spot or with vertex and upper 
frontal area orange. Wings very light brown. Length: 12 mm. -20 mm. 

Head triangular, as wide as or slightly wider than long; tempora promi- 
nent; surface variably sculptured, densely micropunctate, variably punc- 
tate. Pronotum (Fig. 178) nearly quadrate, much wider than long; sides 
nearly straight, divergent from base to near level of apex, then suddenly 
rounded and convergent; disk transversely convex; surface granular, 
densely alutaceous. Elytra weakly to obsolescently rugose. Outer hind 
tibial spur enlarged, obliquely truncate, twice as wide as inner spur; 
inner spur flattened. 

Male. Antennae (Fig. 84) not quite reaching base of pronotum; seg- 
ments IV-X nearly or fully as wide as long; IV-VII weakly incrassate, 
flat on inner side, clothed there with erect pale setae; V and VI, which 
are largest, at most only slightly wider than IX or X. Fore tarsi slightly 
longer than fore tibiae. Emargination of sixth abdominal sternum (Fig. 
242) shallow; lateral lobes of sternum not produced, not hemispherical. 
Pygidium as in Figure 211. Genitalia as in Figure 316; aedeagus with 
ventral hooks subequal, variable. 

Female. Antennae as in Figure 131. 

Type locality. Of tenebrosa, San Diego, California; of occipitalis, 
southern California. 



SYSTEMATICS AND PHYLOGENY 



117 



Geographic distribution. California, from Yolo and Marin counties 
south to San Diego County. ( See Fig. 29. ) 

Seasonal distribution. April 23 — June 15. 

Records. UNITED STATES: California: State label only, 20; Arrow- 
head Springs (Fall, 1901, p. 185); Arvin, March, 4; 20 mi. east of Bakers- 
field (Linsley and MacSwain, 1942); Carmel, May, 1; Clarksburg, April, 
1; Fairfax, May, 1; Gavilan Hills, April, May, 3; Gavilan Peak, Riverside 
County, April, 2; Kern County, 2; Lemoncove, March, 12; Miramar, 
May, 4; Mojave Desert, April, 6; Oil City, Kern River, April, 2; Palomar, 
May, 1; Paraiso Springs, April, 1; Perris, April, 1; Potrero, 3; Riverside, 
April, 6; San Diego, 5 (1 in May); San Diego County, 4 (1 in April); 
Simmler, March, 9; Terminus, Tulare County, March, 4; Visalia, June, 8. 

Remarks. A most unusual feature of this species is the presence of 
erect pale setae on the inner side of some of the intermediate antennal 
segments in the male. A very similar specialization has developed inde- 
pendently in viridana. 

In the present state of our knowledge, this species appears to be 
represented by three morphologically distinct and geographically dis- 




Fig. 29 (left). Distribution of Lytta tenebrosa (circles) and Lijtta lugubris (squares). 
Fig. 30 (right). Distribution of Lytta blaisdelli. 



118 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ltftttt 

continuous populations or races. These intergrade to some extent by 
variation. It should be noted that in comparing the pronotal surfaces of 
the races in the diagnoses given below, reference is made to the basic 
texture of the surfaces exclusive of the alutaceous markings, which are 
about equally strong in all three races. 

The typical race occurs in extreme southern California, from San Diego 
County north to the San Bernardino Mountains. It is characterized as 
follows: head black with a small, rounded or oval orange frontal spot 
which is less than one-third as wide as frontal area between eyes; vertex 
and pronotum finely rugose, strongly granular, dull; vertex moderately 
coarsely, densely punctate; average distance between punctures on vertex 
subequal to diameter of a single puncture; elytra rugose. This is the 
form originally described by LeConte as tenebrosa. 

The northern coastal race occurs from Yolo and Marin counties south 
along the coast to San Luis Obispo County. It differs from the typical 
race as follows: frontal spot larger, triangular or diamond-shaped, at least 
half as wide as frontal area between eyes; vertex smoother, shinier, less 
coarsely, less densely punctate; pronotum slightly smoother; elytra gen- 
erally a little more weakly rugose. In a few specimens the frontal spot 
tends to encroach on the vertex. 

The Central Valley race is found at the southern end of the Central 
Valley of California, in Tulare and Kern counties. In it the pale area 
of the head encompasses both the center of the frontal area (as in the 
northern coastal race) and the vertex, except for a short black median 
stripe. Occasionally there is also a small red mark at the center of the 
pronotal disk. The surface of the vertex is as finely sculptured as in the 
northern coastal race, and the pronotum is even smoother and shinier, 
being nearly as finely sculptured as the vertex. The elytra are obsoles- 
cently rugose. This is the form previously accorded specific rank under 
the name occipitalis. With respect to color, this race intergrades with the 
preceding in that the pale area of the head in a specimen from Oil City 
and in one from Lemon Cove is confined largely to the frontal area. 

The type of tenebrosa is in the Museum of Comparative Zoology. The 
type of occipitalis is in the collection of the Academy of Natural Sciences 
of Philadelphia. 

Bionomics. The specimens from Gavilan Peak were collected from 
Baeria ( Compositae ) . One of the specimens labeled "Mojave Desert" is 
from "sage." 

Some aspects of the life history of this species (as occipitalis) have 
been studied by Linsley and MacSwain (1942). At a nesting site of 
Anthophora linsleyi Timberlake south of Bakersfield, California, these 
authors found tenebrosa infesting 11 or 1.45 per cent of a total of 759 
bee cells examined. According to their observations, the larva may re- 



SYSTEMATICS AND PHYLOGENY 119 

quire the contents of more than one cell for complete development. 
Emergence was recorded in early May. It was noted that adults fed 
primarily on petals of flowers, although some pollen was eaten also. The 
name of the host plant was not given. 

Lytta (Paralytta) blaisdelli (Fall) 

Cantharis blaisdelli Fall, 1909, Canad. Ent., vol. 41, p. 166. 

Lytta blaisdelli, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 83. 

Black. Frontal spot small. Wings pale brown, with at least apical region 
darker and more opaque. Length: 10 mm. -16 mm. 

Head as wide as long; tempora moderately prominent; surface moder- 
ately densely micropunctate, rough to smooth, variably punctate. Pro- 
notum (Fig. 179) as wide as to much wider than long; surface variable. 
Elytra rugose. Hind tibial spurs slender, similar in size; outer spur flat- 
tened or very obliquely truncate; inner spur flattened. 

Male. Antennae (Fig. 85) reaching base of pronotum; segments IV-VII 
moderately incrassate; VII only slightly narrower and shorter than VI, 
obviously longer and wider than IX or X. Fore tarsi as long as fore tibiae. 
Sixth abdominal sternum as in lugubris (see Fig. 243). Pygidium as in 
Figure 212. Genitalia as in Figure 317. 

Female. Antennae as in Figure 132. Fore tarsi slightly shorter than 
fore tibiae. Hind margin of sixth abdominal sternum turned dorsad but 
not abruptly so, not at a right angle to rest of sternum. 

Type locality. Siskiyou County, 2750 ft., California. 

Geographic distribution. A montane species ranging from southwestern 
Idaho to the Klamath Mountains and thence south along the Sierra 
Nevada. ( See Fig. 30. ) 

Seasonal distribution. May 2.5 — July 26. 

Records. UNITED STATES: California: Cantara, Siskiyou County, 
2416 ft.-2750 ft., July, 12; Lake Alpine, July, 2; Paradise Valley, Yosemite 
National Park, July, 1; Siskiyou County, July, 8; Sonora Pass, 9000 ft- 
10,000 ft., July, 135; South Fork of Kings River, 6000 ft., Fresno County, 
May, 1. Idaho: State label only, 4; Boise, June, 2. Nevada: Reno, June, 1. 
Oregon: Garfield Park, Crater Lake National Park, July, 2. 

Remarks. In the series of specimens from Siskiyou County, California, 
the head is relatively smooth and rather finely, sparsely punctate, and the 
wings are pale brown in the anal region basad of vein 1A, becoming 
darker and opaque elsewhere. These characters apply also to specimens 
from the Kings River and Paradise Valley, California. In the specimens 
from Idaho, Nevada, and Oregon the head and pronotum are rough and 
coarsely, moderately densely punctate, the pronotum being, in addition, 
densely alutaceous; the antennal segments of the male are perceptibly 



120 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

less elongate than in the California material mentioned; and the wings 
are dark and opaque only in the apical region. The long series from 
Sonora Pass, California, is like the Idaho, Nevada, and Oregon material 
except that the head and pronotum vary in some specimens to a smooth, 
finely sculptured condition like that of California specimens from other 
localities. A pair of specimens from Lake Alpine, California, are coarsely 
sculptured and otherwise typical of the rough form. 

Coarsely sculptured specimens of blaisdelli closely resemble lugubris. 
These two species are usually separable on the basis of the hind tibial 
spurs, but in some cases it may be necessary to use wing color or char- 
acters of the male genitalia in order to distinguish between them in all 
certainty. 

The first instar larva of blaisdelli has been described by MacSwain 
( 1956 ) . The type is in the Museum of Comparative Zoology. 

Bionomics. Two of the specimens from Sonora Pass, California, are 
recorded from Castilleja; four others from the same locality were taken 
on Pentstemon. Both genera belong to the family Scrophulariaceae. 

Lytta (Paralyita) lugubris (Horn) 

Cantharis lugubris Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 107. 
Schwarz, 1890, Insect life, vol. 2, p. 288. Fall, 1909, Canad. Ent, vol. 41, 
p. 167. 

Lytta lugubris Ulke, 1875, in Report upon the geographical and geological 
explorations and surveys west of the 100th meridian, vol. 5, p. 812, pi. 16, 
fig. 2 (homonymic synonym); 1882, Bull. Brooklyn Ent. Soc., vol. 4, p. 42 
(reprint of 1875 paper). Denier, 1940, Rev. Ent. Argentina, vol. 10, p. 423. 

Cantharis ulkei Beauregard, 1889, Bull. Soc. Ent. France, p. ccxxii (new 
name for lugubris Ulke, unnecessarily proposed); 1890, Les insectes vesicants, 
p. 498 (new name proposed a second time). Fall, 1901, Trans. Amer. Ent. 
Soc, vol. 27, p. 297. 

Black. Frontal spot small, elongate. Wings colorless. Length: 15 mm.- 
19 mm. 

Head triangular, as broad as long; tempora moderately prominent; 
surface rough, densely micropunctate, granular, coarsely, moderately 
densely punctate. Pronotum (Fig. 180) as wide as or slightly wider than 
long; surface as on vertex. Elytra rugose. Hind tibial spurs equal in 
size, moderately thickened, obliquely truncate (Fig. 194). 

Male. Antennae reaching base of pronotum; segments IV-VII moder- 
ately incrassate; VII slightly narrower than VI, obviously longer and 
wider than VIII; VIII noticeably longer and wider than IX or X. Fore 
tarsi as long as fore tibiae. Emargination of sixth abdominal sternum 
(Fig. 243) moderately deep, semicircular; lateral lobes produced, hemi- 
spherical; emargination lined with short, coarse setae. Pygidium as in 
blaisdelli (see Fig. 212). Genitalia as in Figure 318. 



SYSTEMATICS AND PHYLOGENY 121 

Female. Fore tarsi slightly shorter than fore tibiae. Hind margin of 
sixth abdominal sternum turned abruptly dorsad, at a right angle to 
rest of sternum. 

Type locality. Owens Valley, California (Ulke, 1875). 

Geographic distribution. Known from three definite localities, all in 
the Sierra Nevada of California. Recorded also, by state label, from 
Oregon and Nevada. (See Fig. 29.) 

Seasonal distribution. June 27 — July 12. 

Records. UNITED STATES: California: State label only, 6; Hallelujah 
Junction, Lassen County, June, 1; Norvell, July, 8; Cal. [Owens Valley], 
2. Nevada: State label only, 2. Oregon: State label only, 1. Also, from 
the Horn collection, one specimen labeled Washington] T[erritory] 
and three without data. 

Remarks. The specimen from Hallelujah Junction, California, is atypical 
in that the inner hind tibial spur is more obliquely truncate than the 
outer one and the gonostyli of the male genitalia are unusually robust. 

This species has been generally credited to Ulke, but Horn's descrip- 
tion has priority of two years. Lytta higubris Haag-Rutenberg (1880), 
proposed for a species now placed in Epicauta, is a junior homonym of 
Ulke's name (and Horn's), not a senior homonym as Beauregard (1889, 
and 1890) believed when he substituted the name ulkei for higubris 
Ulke. Presumably Beauregard was under the impression that higubris 
Ulke dated from the 1882 reprint of Ulke's description, or, more likely, 
that Haag-Rutenberg's higubris had priority because of its early usage 
( as a nomen nudum ) in the Dejean Catalogue, where it was credited to 
Klug. (Beauregard also erred in giving the range of higubris Ulke as 
Illinois.) The case against ulkei was clearly stated by Denier (1940), 
who proposed for Haag-Rutenberg's higubris the substitute name lugu- 
berrima. 

There are, according to G. Wallace (in litt.), nine specimens under 
the name higubris in the Ulke collection at the Carnegie Museum, each 
bearing the label "Cal." These are undoubtedly the specimens from 
Owens Valley, California, upon which Ulke based his description and 
from which a lectotype of higubris Ulke should be designated. According 
to H. J. Grant, Jr., the Horn collection at the Academy of Natural Sci- 
ences of Philadelphia contains a total of 18 specimens, 2 of which ( a male 
and a female mounted on the same pin) bear the same label as the 
material in the Ulke collection. Because Horn described his higubris from 
specimens received from Ulke, it must be assumed that the pair of speci- 
mens labeled "Cal." in his collection are the only ones eligible for lecto- 
type selection. From this pair I have designated the male as lectotype of 
higubris Horn. 



122 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LljttCl 

Melaena Subgroup 

Black. Head and pronotum smooth, sparsely micropunctate, finely or 
very finely, sparsely punctate. Head quadrate; tempora broadly rounded, 
not prominent; dorsal margin nearly straight. Pronotum campanuliform 
or subcircular, without lateral angles. Scutellum rounded. Elytra rugose. 
Tibiae normally pubescent, lacking lines of setae along inner edges. 
Middle tibiae strongly bowed. Male middle tibiae with a deep, impunc- 
tate, glabrous impression on anterior side apically. Outer hind tibial spur 
about twice as wide as inner spur, obliquely trunctate. Pygidium pro- 
duced. Male genitalia with apical foramen of median tube not extending 
to apex of aedeagus; dorsal sclerite of aedeagus forming a heavy, strongly 
recurved hook. 

Remarks. This subgroup contains the species melaena, morosa, and 
funerea. These are easily distinguished from all other species of Paralytta 
by the strongly modified middle tibiae of the male. The range of the 
subgroup includes the southern two-thirds of California, the Yuma, Ari- 
zona, region, and the peninsula of Baja California. Host plants have been 
recorded only for funerea, which occurs on Compositae. 

Key to Species 

1. Mandibles nearly straight; tarsal pads completely absent, the ventral 
side of tarsi glabrous between ventral fringes; tarsal claws nearly 
straight for basal three-fourths; male sixth abdominal sternum and 
pygidium as in Figures 215 and 246, respectively. . . .funerea (p. 126) 
Mandibles strongly curved; tarsal pads present on at least fore and 
middle legs; tarsal claws curved from base 2 

2. Pronotum ( Fig. 181 ) somewhat quadrate, as wide as long; male 
pygidium ( Fig. 213 ) obtusely rounded or truncate at apex; male sixth 
abdominal sternum with lateral lobes not slender (Fig. 244); female 

fore tarsi nearly as long as fore tibiae melaena (p. 122) 

Pronotum (Fig. 182) subcircular, one-tenth to one-fifth wider than 
long; male pygidium (Fig. 214) very acute at apex; male sixth ab- 
dominal sternum with lateral lobes produced as slender processes 

( Fig. 245 ) ; female fore tarsi four-fifths as long as fore tibiae 

morosa ( p. 124 ) 

Lytta (Paralytta) melaena LeConte 

Lytta melaena LeConte, 1858, Proc. Acad. Nat. Sci. Philadelphia, p. 76. 
Linsley and MacSwain, 1952, Pan-Pacific Ent., vol. 28, p. 134, tbl. Linsley 
and MacSwain, 1958, Wasmann Jour. Biol., vol. 15, pp. 229, 230. 

Cantharis melaena, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 105; 
1896, Proc. California Acad. Sci., ser. 3, vol. 6, p. 371. Fall, 1901, Occas. 



SYSTEM ATICS AND PHYLOGENY 123 

Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc, 
vol. 27, p. 297. 

Black. Frontal spot about one-fourth as wide as frontal area between 
eyes. Apical half of wings brown, basal half almost colorless. Length: 
14 mm.-22 mm. 

Head finely, sparsely punctate. Labium distinctly emarginate. Last 
segment of labial palpi elongate, hatchet-shaped. Pronotum ( Fig. 181 ) 
quadrate campanuliform, as wide as long; disk more coarsely and a little 
more densely punctate than vertex, weakly alutaceous, so as to be duller. 
Hind tarsal pads represented only by some pale setae along ventral edges 
of one or more of second to fourth segments, or completely absent. Tarsal 
claws weakly curved from base. 

Male. Antennae ( Fig. 86 ) with segments IV-VI moderately incrassate; 
VII somewhat compressed but only slightly wider than VIII; VII very 
slightly larger than IX or X. Fore tarsi slightly longer than fore tibiae. 
Pads of fore and middle tarsi well developed, dense, silvery. Emargination 
of sixth abdominal sternum (Fig. 244) moderately deep; lateral lobes of 
sternum moderately produced, rounded. Pygidium (Fig. 213) obtusely 
rounded or broadly truncate at apex. Genitalia as in Figure 319. 

Female. Fore tarsi nearly as long as fore tibiae. Fore and middle tarsal 
pads smaller and less dense than in male. Pygidium rounded. 

Type locality. State of Sonora. 

Geographic distribution. Southern California and the Yuma, Arizona, 
region south to the tip of the peninsula of Baja California. (See Fig. 31.) 

Cockerell's (1898) record of Organ, New Mexico, and Fall's (1901) 
record of Temescal [Alameda County], California, have been rejected 
as probably based on misindentifications. There is a specimen in the 
collection of Cornell University mislabeled Texas. 

Seasonal distribution. March — July 2. 

Records. UNITED STATES: Arizona: Yuma, 1. California: State 
label only, 14 (1 in March); Barrett Springs, April, 3; The Gavilan 
[Peak], March, July, 4; Gilman Hot Springs, May, July, 6; Riverside, 5; 
San Diego, April, May, 5; Tanbark Flat, June, 1; Tehachapi, May, 8. 

MEXICO: Baja California Sur: San Jose de Gracia (Horn, 1896); 
San Jose del Cabo, 2; Santa Margarita Island [Isla Santa Margarita] 
(Horn, 1896). Sonora: (LeConte, 1858). 

Remarks. The specimens from Baja California seem to differ in no 
respect from specimens collected in California. The type is in the Museum 
of Comparative Zoology. 

Bionomics. Larvae and pupae of melaena have been found in resting 
cells in a nesting site of the emphorine bee Diadasia bituberculata 



124 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



Fig. 31. Distribution of Lytta melaena. 

( Cresson ) near Barrett Springs, San Diego County, California ( Linsley 
and MacSwain, 1952 and 1958). 

Lytta (Paralytta) morosa (Fall) 

Cantharis morosa Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 301. 
Lytta morosa, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 82, 
pi. 15. 

Black. Frontal spot about one-third as wide as frontal area between 
eyes. Body frequently in part piceous, especially sides of pronotum. 
Wings light yellowish brown. Length: 12.5 mm. -17.0 mm. 

Head finely, very sparsely punctate. Labium distinctly emarginate. 
Last segment of labial palpi as wide as long, hatchet-shaped. Pronotum 
(Fig. 182) subcircular, one-tenth to one-fifth wider than long; disk con- 
vex; surface as on vertex. Hind tarsal pads represented only by some 
pale setae along ventral edge of second to fourth segments. Tarsal claws 
curved from base. 



SYSTEMATICS AND PHYLOGENY 



125 



Male. Antennae (Fig. 87) with segments IV-VII strongly incrassate; 
VII narrower than VI but obviously longer and wider than VIII; VIII a 
little larger than IX or X. Fore tarsi short, about five-sixths as long as fore 
tibiae. Fore and middle tarsal pads only moderately developed, no larger 
or denser than in female. Emargination of sixth abdominal sternum (Fig. 
245) deeply semicircular, densely lined with short setae; lateral lobes of 
sternum strongly produced, slender, acute. Pygidium (Fig. 214) very 
acute at apex. Genitalia as in Figure 320. 

Female. Antennae as in Figure 133. Fore tarsi about four-fifths as long 
as fore tibiae. Pygidium less acute at apex, much as in male of funerea 
(see Fig. 215). 

Type locality. Temecula, Warners Ranch, or Poway, California. 

Geographic distribution. Extreme southern California, in and near the 
Coastal Range. (See Fig. 32.) 




Fig. 32. Distribution of Lijtta funerea (circles) and Lijtta morosa (squares 



126 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Seasonal distribution. August 27 — September 14. 

Records. UNITED STATES: California: State label only, 3; Alpine, 
September, 5; Gavilan Hills (MacSwain, 1956); Lake Elsinore, Septem- 
ber, 2; Poway, 1; San Bernardino County (Fall, 1901); Temecula, Aug- 
ust, 1 (cotype); Warners Hot Springs, San Diego County, August, 35; 
Warners R[anch], San Diego County, 2. 

Remarks. The first instar larva of this species has been described by 
MacSwain (1956). 

The type of morosa is in the Museum of Comparative Zoology. The 
"cotype" specimen I have examined is in the United States National 
Museum. The specimens from Poway, Temecula, and Warners Ranch are 
probably eutopotypes, but there is no indication that they were included 
in Fall's type series. 

Lijtta (Paralytta) funerea (Fall) 

Cantharis funerea Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 301. 
Lytta funerea, Selander, 1954, Coleopterists' Bull., vol. 8, p. 11. MacSwain, 
1956, Univ. California Publ. Ent., vol. 12, p. 80, pis. 13, 15. 

Black. Frontal spot generally no less than one-third as wide as frontal 
area between eyes. Sides of pronotum, basal half of elytra, and legs 
rarely piceous. Wings very pale yellowish brown. Length: 13.0 mm.- 
18.5 mm. 

Head finely, sparsely punctate. Labrum truncate. Mandibles nearly 
straight, quite unlike those of any other species of the genus. Last segment 
of labial palpi quadrate. Pronotum subcircular, at least as long as wide, 
generally elongate; disk convex; surface as on vertex. Tarsal pads com- 
pletely lacking, the ventral side of segments glabrous between ventral 
fringes. Tarsal claws nearly straight for basal three-fourths, curved at 
apex; ventral blade a little shorter than dorsal one but not fused to it. 

Male. Antennae ( Fig. 88 ) with segments IV-VII moderately incrassate; 
VII shorter and narrower than VI but obviously longer and wider than 
VIII. Fore tarsi as long as fore tibiae. Emargination of sixth abdominal 
sternum (Fig. 246) broadly triangular; lateral lobes of sternum not notice- 
ably produced. Pygidium ( Fig. 215 ) broadly triangular posteriorly. Geni- 
talia as in Figure 321. 

Female. Antennae as in Figure 134. Fore tarsi slightly shorter than fore 
tibiae. Pygidium even more bluntly produced than in male. 

Type locality. Lake, Sacramento, or Tulare County, California. 

Geographic distribution. California, from Riverside County north to 
Santa Cruz, Lake, and (western) Placer counties. (See Fig. 32.) 

Seasonal distribution. March 15 — October 15. 

Records. UNITED STATES: California: State label only, September, 



SYSTEMATICS AND PHYLOGENY 127 

3; Arvin, March, September, 8; Consumnes [River], August, 7; Fresno, 
June, August, September, 26; Indian Wells, Inyo County, May, 2; Kern 
County, March, 3; Kismet and 1 mi. northeast of Kismet, 300 ft., Sep- 
tember, 10; Lake County ( Fall, 1901 ) ; Merced, June, September, 5; 
Merced Falls, September, 1; North Fork, Madera County, October, 2; 
Perkins, September, 49; Ripon, 1; Roseville, August, September, 7; Santa 
Cruz, April, 2; Sequoia National Park, July, 2; Tesla, October, 5; Tulare 
County, 1; western San Joaquin County, 1; Woodlake, 5. 

Remarks. This is the only all-black species of Porolytta lacking tarsal 
pads. With respect to the form of the tarsal claws it approaches moesta 
( Nitidicollis Group ) . 

The first instar larva of funerea has been described by MacSwain 
(1956). The type is in the Museum of Comparative Zoology. 

Bionomics. Adults have been recorded from "tarweed" at Arvin, from 
Hemizonia (Compositae) at and near Kismet, and from Centromadia 
(Compositae) at Woodlake. 

Magister Subgroup 

Usually contrastingly bicolored. Head and pronotum smooth, sparsely 
micropunctate, very finely, weakly, very sparsely punctate. Pronotum 
strongly hexagonal, with prominent, sharp lateral angles. Scutellum 
acutely triangular. Elytra strongly, coarsely reticulate or (some vulne- 
rota) merely rugose. Fore and hind tibiae glabrous on inner side except 
for a fine line of coarse, short setae along each inner edge; middle tibiae 
similar, but with line of setae along posterior inner edge only. Middle 
tibiae strongly bowed. Male middle tibiae more or less concave on 
anterior side apically but lacking a distinct impression. Pygidium pro- 
duced. Male genitalia with apical foramen of median tube not extend- 
ing to apex of aedeagus; dorsal sclerite of aedeagus forming a heavy, 
strongly recurved hook. 

Remarks. This distinctive subgroup contains the species magister, 
vulnerata, and margarita. Its most striking feature is the reticulate condi- 
tion of the elytra, but it should be noted that this character fails in one 
of the races of vulnerata. Elsewhere within the genus Lytta reticulate 
elytra are found only in the Reticulata Group of the present subgenus. 

The Magister Subgroup has a wide distribution in the arid regions of 
North America west of the Rocky Mountains. Species of Compositae 
seem to be the principal host plants of the subgroup, although magister 
has been recorded also from plants of several other families. 

Key to Species 

1. Femora and tibiae orange (rarely piceous, in which case head and 
pronotum are piceous also); head not distinctly triangular; elytra 



128 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

always coarsely reticulate; tarsal pads completely lacking; male sixth 

abdominal sternum as in Figure 247 magister (p. 128) 

Femora and tibiae black; head distinctly triangular; tarsal pads 
present on all but first segment of middle, or middle and hind, tarsi 
2 

2. Vertex (head above eyes) and pronotum orange, with or without a 
median black stripe; elytra varying from coarsely reticulate to rugose; 
emargination of male sixth abdominal sternum deep, rounded (Fig. 

248) vulnerata (p. 130) 

Head (including lower frontal area) and pronotum red; elytra coarsely 
reticulate; emargination of male sixth abdominal sternum shallow, 
very obtusely angulate at apex. Isla Santa Margarita, Baja California 
Sur margarita (p. 136) 

Lytta ( Paralytta ) magister Horn 

Lytta magister Horn, 1870, Trans. Amer. Ent. Soc, vol. 3, p. 90. Tanner, 
1928, Ann. Ent. Soc. America, vol. 21, p. 273. Jaeger, 1938, The California 
deserts, pp. 54-55, fig. MacSwain, 1943, Ann. Ent. Soc. America, vol. 36, 
p. 361, fig. 3; 1956, Univ. California Publ. Ent., vol. 12, p. 79. Selander, 1958, 
Trans. Kansas Acad. Sci., vol. 61, p. 79, fig. 1. 

Cantharis magister, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 106. 
Fall, 1901, Occas. Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. 
Amer. Ent. Soc, vol. 27, p. 296. 

Black. Head, prothorax, femora, and tibiae consistently orange over 
almost the entire range of the species, darkened to piceous black in the 
population of the Pinacate lava cap in extreme northwestern Sonora. 
Some orange or piceous also on epistoma, mandibles, antennal segment 
I, midline of mesothorax and metathorax, coxae, and trochanters; often 
with some orange or piceous on scutellum and sixth abdominal sternum. 
Wings colorless except that apex is a little clouded. Length: 16.5 mm.- 
33.0 mm.; generally more than 20 mm. 

Head slightly wider than long; sides rounded, divergent above eyes; 
head widest below tempora; dorsal margin nearly straight; vertex tumid. 
Pronotum as in Figure 183, one-fifth wider than long; disk flat, fre- 
quently with a shallow, irregular impression on each side between lateral 
angles; center of disk frequently impunctate. Elytra strongly, coarsely 
reticulate. Outer hind tibial spur not much thicker than inner spur, very 
obliquely truncate; truncature elongate. Tarsal pads completely lacking; 
tarsal segments glabrous between ventral fringes. 

Male. Antennae (Fig. 89) reaching at least two segments beyond base 
of pronotum; segments V-VII strongly incrassate. Emargination of sixth 
abdominal sternum (Fig. 247) deeply semicircular, densely lined with 
short, coarse setae; lateral lobes of sternum produced, heavy, broadly 



SYSTEMATICS AND PHYLOGENY 



129 



rounded. Pygidium (Fig. 216) broadly truncate at apex. Genitalia as in 
Figure 322. 

Female. Pygidium truncate, not so strongly produced as in male. 

Type locality. Owens Valley, California. 

Geographic distribution. Lower elevations in southern California, 
southern Arizona, and northwestern Sonora, ranging north along the 
Colorado River and its tributaries to the Grand Canyon and the south- 
western corner of Utah (Washington County). (See Fig. 33.) 

Seasonal distribution. February 28 — October 28. 

Records. UNITED STATES: Arizona: State label only, 2; Bright Angel 
[Creek], 3700 ft., 1; Dripping Springs, Organ Pipe National Monument, 
March, 6; Gila Bend, April, 4; mountain near Gila Bend, February, 3; 
8 mi. south of Hoover Dam, June, 1; Oracle, June, 2; Patagonia, August, 
3; Phoenix, March, April, June, 29; Quartzite, March, 1; Rillito, May, 1; 
Tinajas Altas [Mountains], April, 5; Tucson, April, May, 10; 10 mi. south 
of Tucson, 2000 ft, March, 1; Tucson Mountains, May, 8; Wellton, April, 
1; Winkleman, April, 1; Yuma, 9 (8 in April). California: State label 




Fig. 33. Distribution of Lijtta magister. 



130 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

only, 10; Bagdad, 2; 30 mi. south of Barstow, June, 15; Big Pine, June, 1; 
Borego [Springs], April, 10; Borego State Park, March, 3; Borego Valley, 
April, 4; Carrizo [Creek], March, 2; Death Valley, March, 1; Cronise 
Lake, April, 7; Daggett, April, May, June, October, 11; Furnace Creek, 
Death Valley, April, 1; Hopkins Well, Riverside County, April, 16; Im- 
perial County, March, 16; Independence, May, 2; Indian Wells, Owens 
Valley, May, 8; Indio, April (MacSwain, 1956); 20 mi. east of Indio, 
April, 12; Inyo County, 2; Inyo Mountains, May, 30; Mojave Desert, 2 
(1 in May); Mud Hills, San Diego County, March, 1; Muroc Dry Lake, 
April, 14; Needles, April, 3; Ocotillo, San Diego County, April, 1; Palm 
Canyon, April, 1; Palm Springs, March, April, 33; Panamint Valley, April, 
22; Recordo, June, 1; Rialto, August, 1; Riverside, March, 1; Saltdale, 
May, 1; San Bernardino County, 1; San Diego, 2; Searles Canyon, Lake, 
and Station, April, 78; Witch Creek, 1; Yermo, May, 4. Nevada: State 
label only, 1. Utah: U[tah] Territory], 1; St. George, 10 (1 in May); 
Zion National Park (Tanner, 1928). 

MEXICO: Sonora: Cerro Colorado, ± 35 mi. west-northwest of So- 
noyta, April, 20. 

Remarks. In all specimens of magister from the United States the head, 
pronotum, femora, and tibiae are orange in color. However, in the series 
from the Cerro Colorado crater, Sonora, these areas are darkened to a 
piceous or nearly pure black. Presumably this melanistic variation reflects 
selection for dark coloration against the black background of the Pina- 
cate lava cap of which the Cerro Colorado is a part. Further details are 
given in a separate paper (Selander, 1958). 

The first instar larva of magister has been figured and described by 
MacSwain (1943 and 1956). The type is in the collection of the Academy 
of Natural Sciences of Philadelphia. 

Bionomics. The following host plant associations have been recorded: 
Chrysothamnus paniculatus (Compositae) at Zion National Park, Utah 
(Tanner, 1928); Coldenia palmeri ( Boraginaceae ) (determined by R. 
McVaugh) at Cerro Colorado crater, Sonora; Geraea canescens (Com- 
positae) at Hopkins Well, California; Lupinus ( Leguminosae ) and un- 
specified Compositae at Muroc Dry Lake, California; lupine, 30 mi. south 
of Barstow, California; and yucca (Liliaceae) at Oracle, Arizona. 

According to the account given by Jaeger (1938), magister at times 
occurs in such numbers in the deserts of California that the scraping of 
the beetles' elytra against stones and dead vegetation fills the air with 
a rasping sound. 

Lytta (Paralytta) vulnerata (LeConte) 

Cantharis vulnerata LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 159. Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. 106. Saunders, 1876, 



SYSTEMATICS AND PHYLOGENY 131 

Canad. Ent, vol. 8, p. 224, fig. 10. Champion, 1892, Biol. Centrali-Americana, 
Coleoptera, vol. 4, pt. 2, p. 437. Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, 
p. 296; 1901, Occas. Papers California Acad. Sci., vol. 8, p. 185. 

Lytta vulnerata, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 331. Horn, 1867, Proc. Ent. Soc. Philadelphia, p. 292; 1870, Trans. Amer. 
Ent. Soc, vol. 3, p. 90 ; 1872, in Rept. U.S. Geol. Surv. 1871, 1872, p. 390. 
Ulke, 1875, in Report upon the geographical and geological explorations and 
surveys west of the 100th meridian, vol. 5, p. 825. Knowlton, 1930, Florida 
Ent., vol. 14, p. 53. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, 
p. 81, pi. 14. 

Lytta coo-peri LeConte, 1854, Proc. Acad. Nat. Sci. Philadelphia, vol. 7, 
p. 18; 1857, in Reports of explorations and surveys . . . from the Mississippi 
River to the Pacific Ocean, vol. 12, book 2, pt. 3, p. 51, pi. 2, fig. 6. Horn, 

1870, Trans. Amer. Ent. Soc, vol. 3, p. 90 ; 1872, in Rept. U.S. Geol. Surv. 

1871, 1872, p. 390. Knowlton, 1939, Utah Agr. Exp. Sta. Mimeo. Ser. 200 
(Tech.), pt. 3, p. 5. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, 
p. 82. 

Cantharis vulnerata var. cooperi, Horn, 1873, Proc. Amer. Philos. Soc, 
vol. 13, p. 106. Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 296. 

Cantharis anguicollis Duges, 1889, An. Mus. Michoacano, vol. 2, p. 105. 

Cantharis cooperi, Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 304. 

Lytta vulnerata cooperi, Knowlton and Taylor, 1952, Utah Agr. Exp. Sta. 
Mimeo. Ser. 389, p. 15. Woodbury, 1956, Ecological check lists. The Great 
Salt Lake Desert series, p. 37. 

Black. Vertex orange, often with a median black stripe or fine line; 
head rarely all black except for a small orange frontal spot. Pronotum 
orange with a black or piceous anterior marginal area, or with black 
area extending posteriad as a median stripe. Wings colorless except that 
apex is clouded. Length: 9 mm. -27 mm. 

Head strongly triangular, as wide as to one-fifth wider than long; 
tempora prominent; dorsal margin varying from nearly straight to deeply 
concave. Pronotum as in Figure 184, one-fifth to one-fourth wider than 
long; disk flat at center, becoming rather sharply declivous at sides 
before and behind lateral angles, almost always with a shallow fovea 
or irregular impression on each side between lateral angles; center of 
disk frequently impunctate. Elytra varying from coarsely reticulate to 
rugose. Outer hind tibial spur at most twice as wide as inner spur, 
obliquely truncate. Fore and middle tarsal pads poorly developed, those 
of at least some segments divided on midline; hind tarsal pads represented 
only by some pale pubescence along ventral edges of second to fourth 
segments. 

Male. Antennae (Fig. 90) reaching base of pronotum; segments IV-VII 
moderately incrassate; VIII slightly so, not compressed. Emargination 
of sixth abdominal sternum (Fig. 248) deep, rounded; lateral lobes of 
sternum strongly produced, rounded and densely pubescent at apex. 
Pygidium ( Fig. 217 ) subacute at apex. Genitalia as in Figure 323. 



132 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Female. Antennae as in Figure 135. First segment of middle tarsi lack- 
ing pad (pale pubescence). Pygidium obtuse at apex. 

Type locality. Of vulnerata, San Diego, California; of cooperi, "Wenass 
River to Fort Colville" [Wenas Creek to Colville, Washington]; of an- 
guicollis, Baja California. 

Geographic distribution. Intermontane plateaus of western North 
America from southern British Columbia south through the Columbia 
Plateau, the Great Basin, and the Colorado Plateau region of Utah to 
the Mojave Desert; thence southwest to the coastal ranges of California, 
extending north ( west of the Sierra Nevada ) to Sacramento County, Cali- 
fornia, and south onto the peninsula of Baja California. (See Fig. 34.) 

Cockerell's (1898) record (also listed by Fall and Cockerell, 1907) of 
vulnerata from the Mesilla Valley, New Mexico, and Gibson's (1912) 
record of cooperi from Indian Head, Saskatchewan, are unacceptable on 
geographic grounds. The latter is based on a series of seven specimens 
in the Canadian National Collection labeled "Indian Head, A. P. Saun- 
ders." 



Fie. 34. Distribution of Lijtta vulnerata. 



SYSTEMATICS AND PHYLOGENY 133 

Seasonal distribution. April 21 — December 30 (also one record in 
February). Collected most commonly from August to October. 

Records. CANADA: British Columbia: Vernon, 1. 

UNITED STATES: Arizona: State label only, 1. California: State 
label only, 12; Antioch, August, September, 55; Azuza, 3 (2 in July); 
Bakersfield, September, 6; Bodfish, 3; 2 mi. east of Caliente Mountain, 
October, 7; Campo, San Diego County, September, 2; Cantua Creek, 
November, 1; Castaic, February, 3; Claremont, September, 15; Claremont, 
mountains near, 4; Coachella Valley, November, 1; Coalinga, 9 (4 in 
September); Colton, September, October, 14; Corona, 3 (2 in Septem- 
ber); Cuyama Valley, Ventura County, October, 2; Desert Spring, 7 mi. 
north of San Bernardino, August, 40; Fillmore, September, 5; Fort Tejon 
(Horn, 1873); Fresno, 2; Garnet, December, 1; Hallelujah Junction, Las- 
sen County, July, August, 47; Hollister, August, 2; Indio, August, 1; Kings 
County, September, 6; Lake Elsinore, September, 1; Lebec, September, 5; 
Littlerock, October, 15; Los Angeles County, 1; Mill Creek Canyon, San 
Bernardino County, September, 7; Mojave, 9 (7 in October); Mount 
Jupiter, 3100 ft. (not located), August, 3; Ontario, September, 1; Owens 
Valley (Horn, 1873); Palmdale, October, 51; Palm Springs, April, July, 
October, 3; Panamint Mountains, October, 1; Pasadena, August, Septem- 
ber, 5; Playa del Ray, Los Angeles County, October, 11; Pomona (Fall, 
1901, p. 185); Sacramento, August, 1; San Bernardino, August, October, 
7; San Bernardino County, September, October, 4; San Diego, 2; San 
Diego County, 1; San Felipe Valley, San Diego County, October, 3; 
S[anta] Maria River, September, 3; Shafter, October, 1; Tehachapi, Aug- 
ust, 7; Tesla, September, 1; Warners Hot Springs, August, 3; White Water, 
September, October, 4. Idaho: Blackfoot, August, 6; Bliss, September, 1; 
Boise, 1; Camas Creek (ambiguous, not mapped), 6; 4 mi. north of 
Hazelton, September, 2; Hunt Project, 2 mi. north of Hazelton, July, 1; 
Idaho Falls, September, 1; Indian Massacre Rock (not located), August, 
1; Jordan River, Owyhee County (not located), 1; Juniper Creek, Owyhee 
County, August, 1; Notus, August, 1; Parma, 2224 ft., September, 2; Poco- 
tello, 16 (5 in August); Ross' Fork, August, 3; St. Anthony, 4956 ft., 
August, 8. Nevada: State label only, 12; Currant Creek, Nye County, 
July, 7; Deeth, July, 3; Las Vegas, September, October, 11; Pyramid Lake, 
August, 16; Ruby Mountains, 1; Volma (not located), July, 1; Wadsworth, 
September, 21; Walker Lake, September, 56. Oregon: State label only, 5; 
Arlington, September, 9; Freewater, September, 1; lone, September, 2; 
Island City, 1; Ontario, September, 3; Paisley, August, 1. Utah: 2 mi. east 
of Arlington Drive, [Salt Lake City], September, 2; Blue Creek, Box 
Elder County, August, 2; Cache Junction, Cache County, August, 27; 
Delta, August, 2; Desert Range Experiment Station, 50 mi. west of Mil- 
ford, 5; Duchesne, July, August, 15; Dugway Proving Grounds, Tooele 



134 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytt(l 

County, August, September, 36; Emery County, October, 2; F[or]t Du- 
chesne, Uintah County (Knowlton, 1939); Hanksville, September, 1; 
Hansel Mountains, August, 1; Howell, August, 8; Lampo, Box Elder 
County (Knowlton, 1930); Lapoint, 5500 ft., August, 1; Little Granite 
Mountain, Tooele County, August, 6; Logan (Knowlton, 1930); Manila 
(Knowlton and Taylor, 1952); Manti, August, 1; Milford, August, 3; 
Millard [County], August, 5; Myton, August, 2; Roosevelt, June, August, 
3; St. George, 1; 2 mi. north of San Rafael [River] bridge, Emery County, 
October, 7; Watson, Uintah County, June, 1; Waw Waw Springs, Beaver 
County, September, 1. Washington: Coulee City, August, 1; Fort Colville 
[Colville] (LeConte, 1857); Kahlotus, August, 6; 5 mi. south of Lowden, 
August, 2; Prosser, June, August, October, 3; Selah, 1150 ft., September, 
4; Stratford, September, 22; Walla Walla, September, 10; Wenass River 
[Wenas Creek] (LeConte, 1857); Yakima, August, 10. 
MEXICO: Baja California Norte?: Baja (or Lower) California, 5. 

Remarks. Adults of this species vary geographically in characters of 
elytral sculpturing, color, body size, and shape of the pronotum. This 
variation is described in the following paragraphs. 

In all specimens from Baja California, the coastal ranges of California, 
and the northern and central parts of the Central Valley of California 
both the head and pronotum have a rather broad black stripe medianly. 
The pronotal stripe tends to be weakened or broken in the basal half in 
some of these specimens but is complete in the majority. The series from 
Playa del Ray, California, has particularly heavy markings: the vertex is 
entirely black, and the stripe on the pronotum is broadened to cover all 
but the lateral angles. On and near the western and northern limits of 
the Mojave Desert in California there is a well-marked tendency for the 
head stripe to be narrowed to a fine median line and the pronotal stripe 
to be broken in the basal half. In occasional specimens from this area 
the head stripe is absent, and the pronotal stripe is limited to the anterior 
half (or less) of the pronotum. Eastward, through the rest of the range 
of the species, the vertex of the head is entirely orange, except in very 
rare instances in which a fine median line is present. In this same region 
the median pronotal stripe is invariably reduced to a small anterior 
marginal area. 

Although additional records are needed to precisely establish the 
geographic pattern of the color variation, it now appears that the reduc- 
tion of the black stripe of the head and pronotum takes the form of a true 
cline, beginning on the western edge of the Mojave Desert and extending 
along its northern edge as far east as Las Vegas, Nevada. 

Variation in the sculpturing of the elytra shows excellent concordance 
with the variation in color just discussed. Thus in Baja California and 
western California the elytra are invariably rugose. Along the edges 



SYSTEM ATICS AND PHYLOGENY 135 

of the Mojave Desert there is a slight tendency for development of 
reticulations. In the specimen from the Panamint Mountains, California, 
and in the series of six specimens from Las Vegas, Nevada, a regular 
system of weak reticulations is evident. Finally, in all specimens from 
the Great Basin and areas to the north and east the elytra are as strongly 
reticulate as in m agister. 

Geographic variation in body size is shown by the fact that the 
average length of 73 specimens from California and Baja California is 
18.6 (10-27) mm., as compared with an average of 15.3 (9-22) mm. for 
a series of 133 specimens from Nevada, Utah, Idaho, and Washington. I 
have not attempted to pursue the study of size variation any further. 

With respect to variation in the shape of the pronotum, there seems 
to be a tendency for the lateral angles to be somewhat less acute in speci- 
mens from California and Baja California than in those from other areas. 
Again, I have not studied this particular variation in detail. 

Geographic variation in the larvae of vulnerata has been described 
by MacSwain (1956), who found minor differences between specimens 
from coastal California (Antioch, Contra Costa County) and specimens 
from the western edge of the Great Basin (Hallelujah Junction, Lassen 
County, California). MacSwain treated the two series under the names 
vulnerata and cooped, respectively, but indicated that he did not con- 
sider the larval differences to be of a specific nature. 

In view of the above discussion, and in particular the high degree of 
concordance shown between variation in color and elytral sculpturing, 
it seems convenient to continue the practice of making a distinction be- 
tween the Baja California and California population (typical vulnerata) 
and the population occurring east of the Cascade-Sierra Nevada range 
( cooperi ) . As names for these populations I suggest "California race" and 
"interior race," respectively. 

In one of the interior race males from Pyramid Lake, Nevada, the 
lateral margins of the pronotal disk are strongly spinose. Four spines are 
present on the left side, two of these at the lateral angle and two along 
the posterolateral margin. These spines increase in length posteriorly, 
the fourth or last one being longer than antennal segment II. The right 
side of the pronotum is similarly modified except that there is no spine 
corresponding to the first spine of the left side and that the anterior 
two spines present are subequal in length. In addition to this male, there 
is a female from the same locality and two females from nearby Halle- 
lujah Junction, California, in which rudiments of two spines are present 
on each of the lateral angles of the pronotum. Under the circumstances 
it seems reasonable to consider this anomaly as genetically controlled. 

In calling attention to this pronotal anomaly in vulnerata, it is worth 
noting that Fall's description of the pronotum of the species margarita, 



136 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytttt 

which was based on a unique male from Isla Santa Margarita, off the 
coast of the peninsula of Baja California, seems to agree very closely 
with the condition of the Pyramid Lake male. This similarity is possibly 
a fortuitous one, but inasmuch as vulnerata appears to be closely 
related to margarita, it is tempting to speculate that the spinose condi- 
tion is genetically homologous in the two species. 

The types of vulnerata and cooperi are in the Museum of Comparative 
Zoology. The type of anguicollis has been lost. There is, however, a speci- 
men from "Baja California" in the Duges collection at the Instituto de 
Biologia in Mexico City that probably belonged to the series from which 
Duges described anguicollis; I have designated this specimen as neotype. 

Bionomics. Specimens have been collected on the following Compositae: 
Artemesia (California), Baccharis (California; LeConte, 1853), Chryso- 
thamnus (California and Nevada), C. nauseosus and C. stenophyllns 
(Utah), Gutierrezia (California and Idaho), and Solidago (Washington). 

Lytta (Faralytta) margarita (Fall) 
Cantharis margarita Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 300. 

According to Fall's description, margarita closely resembles the interior 
race of vulnerata. The main differences between the two are that in 
margarita the color of the head and prothorax is red (rather than orange), 
the lower frontal area of the head is not black, the intermediate seg- 
ments of the male antennae are "scarcely perceptibly thickened," the 
tibiae are "much stouter" and "less compressed," and the emargination of 
the male sixth abdominal sternum is shallow, with the apex very obtusely 
angulate. The pronotum was described as having "three acute tubercles 
along the side margin posteriorly." This was evidently a bilaterally sym- 
metrical condition as in his key to species Fall described the pronotum as 
"trituberculate each side behind the lateral angles." A similar condition 
occurs as a rare anomaly in the interior race of vulnerata, as I have 
already noted. 

Type locality. Isla Santa Margarita, Baja California Sur. 
Geographic distribution. Known onlv from the type locality. (See Fig. 
36. ) 

Seasonal distribution. Unknown. 

Records. MEXICO: Baja California Sur: Isla Santa Margarita (Fall, 
1901). 

Remarks. There are apparently no specimens of this species in collec- 
tions at the present time. The original description was based on a unique 
male in the collection of Charles Fuchs. If this specimen were still in 
existence, it would probably be in the collection of the California Acad- 



SYSTEM ATICS AND PHYLOGENY 137 

emy of Sciences, but Hugh B. Leech has informed me (in lift.) that it is 
not there. In all probability it was lost in the San Francisco fire, which 
destroyed the greater part of the Fuchs collection. 

Subgenus ADICOLYTTA, new subgenus 

Moderately large, elongate beetles. Colors nonmetallic; head when 
black with a pale frontal spot. Upper surface essentially glabrous. 
Head (at least vertex) and pronotum finely granular, micropunctate, 
finely, sparsely, or very sparsely punctate. Eyes rather narrow, not promi- 
nent. Male antennae slightly longer than in female, just short of base 
of pronotum or extending one or two segments beyond; male inter- 
mediate antennal segments (at least IV and V) incrassate, wider than 
following segments; except in erebea, two or more of the intermediate 
segments with conspicuous, asymmetrical distortion (antennae serrate 
in peninsularis) . Female antennae clavate. Pronotum with sides rounded 
and gradually convergent from middle to base, more abruptly convergent 
to apex; disk lacking calluses. Elytral surface rugose or rugose punctate, 
granular. Except in peninsularis, male trochanters with modified pubes- 
cence, male middle tibiae and either fore tibiae or middle femora vari- 
ously modified, and first segment of male middle tarsi distorted. Each 
tibia with two spurs. Hind tibial spurs strongly dissimilar; outer hind 
tibial spur two to three times as wide as inner spur, obliquely truncate; 
inner spur only slightly thickened, excavate. Tarsal claws rather heavy, 
moderately curved. Pygidium not produced. Male genitalia with gono- 
styli lacking mesal hooks, either slender, tapered, and evenly rounded 
at apex or somewhat heavier and weakly lobed laterally before apex; 
minute, sparse setae present on gonostyli; aedeagus with two well- 
developed ventral hooks. 

Type species. Cantharis mutilata Horn. 

Remarks. This subgenus contains seven well-marked species, divided 
into three groups. It is limited in distribution to the North American 
Continent and within this region occurs primarily in Mexico, one species 
(mutilata) ranging north into the southwestern United States and another 
( eucera ) south into western Panama ( Fig. 6 ) . It most closely resembles 
the subgenus Paralytta, and, as pointed out above, it seems to be most 
closely related to that subgenus. Although it is readily diagnosed by a 
combination of characters, there seems to be no single character that 
will distinguish all members of the subgenus from those of all other sub- 
genera of Lytta. With some experience, however, it is easily separable 
on the basis of the general facies of the adults. 

In at least eucera, sanguinca. erythrothorax, and mutilata, and prob- 
ably in the other species of the subgenus as well, the abdomen in life 



138 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

is swollen to the extent that the membranous areas of the third to sixth 
terga are partially exposed at the sides of the elytra. These areas are 
bright red or orange in color and as a result are very conspicuous in 
those species in which the elytra and abdomen are black. Possibly they 
are aposematic in function. This does not hold for sanguinea, where the 
membranous areas are about the same color as the rest of the beetle. 

Very little is known of the bionomics of the species of the subgenus. 
There is, however, enough data available to suggest that the Eucera and 
Mutilata groups may be separable on the basis of host plant differences, 
the former being perhaps restricted to Cucurbitaceae and the latter to 
Convolvulaceae. There are no host plant records for the Peninsularis 
Group. 

The relationships of the species of the subgenus Adicolytta seem 
reasonably clear. The postulated phylogeny is shown in the diagram in 
Figure 35. The same arrangement has been followed in constructing the 
kevs to groups and species. 

In arriving at the arrangement indicated, I have assumed that the 
following characters are primitive for the subgenus: antennae moniliform, 
with segments V and VI incrassate in male but not distorted; male legs 
lacking obvious sexual modification (as in peninsularis); tarsal pads 
only moderately developed in both sexes (as in peninsularis); and male 
genitalia with gonostyli slender, tapered, and evenly rounded at apex 
(as in the Peninsularis and Eucera groups). This combination of char- 
acters presumably once existed in an ancestral form. 

From such an ancestral type the species peninsularis may be derived 
separately as a line in which the antennae became serrate in the male 
( subserrate in the female ) . At present peninsularis seems to be restricted 
in range to the southern tip of the peninsula of Baja California. However, 
it seems likely that at one time the line to which it belongs was more 
widely distributed, probably extending onto the Mexican mainland, and 
that subsequently it was eliminated over most of its range. 

The Eucera and Mutilata groups apparently arose from an ancestor in 
which antennal segments V and VI were weakly expanded on the dorsal 
side (as in erebea), the tarsal pads were enlarged and dense, and the male 
legs had developed specializations, at least to the extent that the pubes- 
cence of the trochanters was somewhat modified and the middle femora 
and the first segment of the middle tarsi were distorted. The Eucera 
Group is more primitive than the Mutilata Group in having male genitalia 
essentially like those of peninsularis. Presumed specializations in the 
former group include the asymmetrical distortion of the hind tarsi and 
the peculiar compression and distortion of the male fore and middle 
tibiae. In all characters except possibly color erebea seems to be the 
more primitive of the two species constituting the group. For example, 



SYSTEMATICS AND PHYLOGENY 



139 



in erebea male antennal segments IV-VI are only weakly expanded 
dorsally, while in eucera segment IV is strongly expanded and V and VI 
are very strongly produced. Again, in erebea the male middle tibiae, while 
contorted and excavate much as in eucera, lack the modification of the 
pubescence found in that species. Interestingly, erebea has a small geo- 
graphic range and seems to be rare, while eucera has the largest range 
of any species of the subgenus and would appear to be the most success- 
ful. 

It is presumed that from a common ancestor with the Eucera Group a 
second line developed which gave rise to the Mutilata Group. In this 
line male antennal segments V and VI were more markedly produced 
dorsally, the male middle femora developed a concavity on the anterior 



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Fig. 35. Phylogeny of the subgenus Adicolytta. 



140 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

side, the distortion of the first segment of the male middle tarsi was in- 
tensified, and the gonostyli of the male genitalia became heavier and 
developed a weak lobe laterally. Within the Mutilata Group sanguined 
appears to be the most primitive species structurally. However, its 
entirely pale color is atypical of the subgenus, and I believe on this 
account that it is to be regarded as a specialized character. In the branch 
of the line leading to erythrothorax, mutilata, and cardinalis male 
antennal segment V became strongly produced, and the male middle 
femora became more highly modified than in the common ancestor with 
sanguined. The species erythrothorax apparently evolved without fur- 
ther major change, while mutilata and cardinalis were derived from a 
separate offshoot in which male antennal segment V became even more 
strongly distorted than in erythrothorax. There is a difficulty with this 
hypothesis in that sanguinea and erythrothorax have the male fore femora 
modified on the ventral side, while mutilata and cardinalis do not. It is 
therefore necessary to assume that either the modification existed at the 
beginning of the Mutilata Group line and was subsequently lost in the 
common ancestor of mutilata and cardinalis or, perhaps equally likely, 
that it is of independent origin in sanguinea and erythrothorax. Attempts 
to resolve this difficulty by arranging the species differently in the phy- 
logenetic tree only lead to greater difficulty in accounting for the distri- 
bution of other characters within the group. 

Key to Groups 

1. Antennal segments IV-X triangular (males, Fig. 91) or subtriangular 
( females ) , produced on ventral side apically; under surface of thorax 
densely clothed with long, shaggy pubescence; tarsal pads only 
moderately developed, tending to be divided on midline; male legs 
unmodified; black with reddish head and pronotum. Cabo San Lucas, 

Baja California Sur Peninsularis Group (p. 141) 

At least distal antennal segments moniliform, nearly symmetrical (IV 
and following one to three segments expanded or produced on dorsal 
side in male); under surface of thorax sparsely clothed with short 
pubescence; tarsal pads well developed, very dense, cushion-like, 
undivided; male legs variously modified 2 

2. Pronotum and elytra black; male middle tibiae contorted, excavate; 
first segment of male middle tarsi normally, sparsely pubescent; first 
three hind tarsal segments clothed laterally with long, hairlike setae; 

pads of these segments asymmetrical Eucera Group ( p. 143 ) 

Color not as above; male middle tibiae modified but not contorted 

or excavate; hind tarsal segments unmodified 

Mutilata Group ( p. 147 ) 



SYSTEMATICS AND PHYLOGENY 141 

PENINSULARIS GROUP 

Black with red head and pronotum. Male antennae serrate; segments 
IV-X triangular, produced on ventral side apically. Female antennae 
subserrate; segments IV-X subtriangular. Scutellum with fine punctures 
and setae. Under surface of thorax, coxae, trochanters, and base of femora 
densely clothed with long, shaggy, erect pubescence; many of the setae 
curved. Male legs unmodified. Tarsal pads only moderately developed, 
not cushion-like, those of all but first segment of each tarsus divided or 
parted on midline; hind tarsal pads smaller and more widely divided than 
rest. Male fifth abdominal sternum evenly, moderately deeply emarginate. 
Female sixth sternum shallowly, subtriangularly emarginate. Genitalia 
with gonostyli slender, tapered, evenly rounded at apex, as in Eucera 
Group except that setae are shorter and sparser; apex of aedeagus drawn 
out to a blunt peak; dorsal hook slender, not strongly recurved. 

Remarks. This group contains only peninsular is, which is known from 
a single locality on the southern tip of the peninsula of Baja California. 

Lytta (Adicolytta) peninsularis (Fall) 
Cantharis peninsularis Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 300. 

Black. Vertex and upper frontal area of head reddish orange. Pro- 
thorax reddish orange; pronotum margined with black, very finely so 
on anterior half. Wings colorless, except for a yellow cast at apex; veins 
of costal margin brown, rest yellow. Length: 18 mm. -20 mm. (Fall, 1901 ). 

Head a little longer than wide; sides subparallel above eyes; tempora 
smoothly rounded; dorsal margin weakly rounded; vertex finely alutace- 
ous, moderately shiny, sparsely punctate; frontal area similar but moder- 
ately densely punctate, three shallow impressions between eyes, lateral 
impressions deeply, longitudinally alutaceous. Maxillary and labial palpi 
slender. Pronotum with distinct lateral angles, as wide as long, widest at 
middle; disk flattened before middle, a wide, shallow impression on each 
side behind middle; surface smoother than on vertex, finely, very sparsely 
punctate at center, a little more densely punctate at sides. Elytra rugose, 
subimpunctate. Tarsi moderately slender. 

Male. Antennae ( Fig. 91 ) serrate, compressed, reaching two segments 
beyond base of pronotum; segments III-X progressively shorter, except 
that V and VI are equal in length; IV-X triangular, distinctly arcuate 
on ventral side, produced on ventral side at apex. Middle tibiae bowed. 
Pad of first segment of hind tarsi absent. Emargination of sixth abdominal 
sternum (Fig. 249) very deeply, acutely triangular; sternum with a pair 
of obtuse ridges diverging from apex of emargination. Genitalia as in 
Figure 324. 



142 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 



Female. Antennae stouter; segments IV-VII subtriangular. Sixth ab- 
dominal sternum shallowly, subtriangularly emarginate. (Female char- 
acters from Fall, 1901.) 

Type locality. Cabo San Lucas, Baja California Sur. 

Geographic distribution. Known only from the type locality. ( See Fig. 
36. ) 

Seasonal distribution. Unknown. 

Records. MEXICO: Baja California Sur: Cape [Cabo] San Lucas, 1. 

Remarks. The unusual antennae, hairy under surface, uniquely modi- 
fied sixth abdominal sternum of the male, and absence of sexual modifica- 
tion of the legs of the male are among the more obvious diagnostic 
characters of this distinct species. Superficially it resembles mutilata, 
which has essentially the same color pattern. 

The type of peninsularis is in the Museum of Comparative Zoology. 
The specimen I have studied is a male from the Van Dyke collection in 
the California Academy of Sciences. It is probably a eutopotype, al- 
though it does not bear Fall's label. 




Fig. 36. Distribution of Lijtta peninsularis (circle), Ltjtta erebea (squares), and Lytta 
margarita ( triangle ) . 



SYSTEMATICS AND PHYLOGENY 143 

EUCERA CROUP 

Pronotum and elytra black. Male antennal segments IV-VII enlarged, 
either expanded or produced on dorsal side. Female antennae slender, 
more strongly clavate than in Mutilata Group. Scutellum impunctate, 
glabrous. Under surface sparsely clothed with short pubescence. Legs 
slender. Setae of male fore and middle trochanters long, dense, erect 
ventrally. Male fore tibiae moderately compressed, enlarged apically, 
slightly twisted. Male middle tibiae strongly compressed, enlarged api- 
cally, strongly contorted; mesal surface excavate. Male fore and middle 
tarsi slightly expanded in relation to those of female. First segment of 
male middle tarsi weakly distorted, sparsely pubescent. Tarsal pads well 
developed, very dense, cushion-like; clothing setae on outer side of first 
three hind tarsal segments long, hairlike, encroaching to a greater or 
lesser extent on pads of these segments, which are asymmetrical. Male 
fifth abdominal sternum entire or nearly so. Male sixth sternum impunc- 
tate, glabrous, and flat at middle posteriorly, moderately deeply emargi- 
nate. Female sixth sternum subtruncate. Male genitalia with gonostyli 
as in Peninsularis Group; apex of aedeagus bluntly rounded; dorsal hook 
strongly recurved. 

Remarks. This group contains the species erebea and eucera. 

Key to Species 

1. Head black with a small orange frontal spot; elytra distinctly rugose; 
segments V and VI of male antennae not strongly produced on one 

side (Fig. 92) erebea (p. 143) 

Head with at least tempora and a large frontal spot orange; elytra 
obsolescently rugose; segments V and VI of male antennae very 
strongly produced on one side ( Fig. 93 ) eucera ( p. 144 ) 

Lytta (Adicolytta) erebea (Champion) 

Cantharis erebea Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 442, pi. 20, fig. 20. 

Black. Head with a small orange frontal spot. Wings dark brown. 
Length: 20 mm. -23 mm. 

Differs structurally from eucera as follows. 

Head as wide as long, not at all, or not nearly as distinctly, triangular 
in form; tempora more evenly rounded. Surface of head and pronotum 
more strongly granular, more densely, strongly micropunctate, less shiny 
(only as shiny as in erythrothorax) , a little less finely punctate. Pronotum 
not as definitely hexagonal, as long as wide; disk more convex, a wide 
shallow fovea on each side behind middle. Elytra more coarsely, strongly 
rugose (as in erythrothorax). 



144 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Male. Antennae (Fig. 92) shorter, reaching base of pronotum; segment 
I short, bulbous; IV-VI moderately incrassate, weakly expanded on dorsal 
side and therefore a little more strongly bulged on that side than on 
ventral side; V largest; VII very slightly enlarged, intermediate in size 
and shape between VI and distal segments. Middle tibiae with excavate 
apical area normally pubescent. Genitalia as in Figure 325. 

Female. Antennae a little more gradually clavate than in eucera. 

Type locality. Huetamo, Michoacan. 

Geographic distribution. Southeastern Michoacan and southern Mexico 
(state). Both localities recorded for the species are within the drainage 
system of the Rio Balsas. (See Fig. 36.) 

Seasonal distribution. Unknown. 

Records. No data, 1. MEXICO: Mexico: State label only, 1 (paratype); 
Temescaltepec, 2. Michoacan: Huetamo, 1 (paratype). 

Remarks. Judged by its representation in collections, erebea is one of 
the rarest Mexican species of Lytta. It was described originally from 
a male and three females, now in the British Museum ( Natural History ) . 
I have studied two of the females from the type series; a male without 
data in the Duges collection in the Instituto de Biologia, Mexico, D. F.; 
and a male and female collected at Temescaltepec in 1931 by G. B. Hinton. 

Figures of the genitalia and the antennae of the type male were 
kindly furnished me by C. M. F. von Hayek. The specimen figured differs 
from the type in that the anterior end of the aedeagus is more strongly 
bent. 

Lytta (Adicolytta) eucera (Chevrolat) 

Cantharis eucera Chevrolat, 1834, Coleopteres du Mexique, fasc. 3, p. [19]. 
Herrera and Mendoza, 1866, Gaceta Med. Mexico, vol. 2, p. 265. Duges, 
1869, La Naturaleza, vol. 1, p. 100, pi. 1, fig. 9. Horn, 1873, Proc. Amer. 
Philos. Soc, vol. 13, p. 104. Duges, 1889, An. Mus. Michoacano, vol. 2, 
p. 101. Champion, 1892, Biol. Centrali-Amerieana, Coleoptera, vol. 4, pt. 2, 
p. 438. Fall, 1901, Trans. Amer. Ent. Soc., vol. 27, p. 296. 

Cantharis spectabilis Castelnau, 1840, Histoire naturelle des insectes Cole- 
opteres, vol. 2, p. 273. Lacordaire, 1859, Genera des Coleopteres, vol. 5, p. 
676. Haag-Rutenberg, 1879, Deutsche Ent. Zeitschr., vol. 23, p. 412. 

Cantharis encera [sic], Horn, 1883, Trans. Amer. Ent. Soe., vol. 10, p. 310 
(lapsus calami) . 

Lytta eucera, Denier, 1940, Rev. Ent., vol. 11, p. 802. Dillon, 1952, Amer. 
Midland Nat., vol. 48, p. 348. Selander, 1959, Proe. Ent. Soe. Washington, 
vol. 61, p. 205, fig. 1. 

Black. Vertex and upper frontal area dark orange or red except for 
a wide longitudinal median band, or only tempora and a heart-shaped 
frontal spot red or orange. Wings dark brown. Length: 14 mm. -32 mm. 

Head wider than long; sides strongly divergent above eyes; tempora 



SYSTEMATICS AND PHYLOGENY 145 

strongly rounded, prominent; dorsal margin straight, a shallow notch 
on midline; surface smooth, very shiny, only faintly granular, very sparsely 
punctate. Pronotum distinctly hexagonal, a little wider than long, widest 
before middle; disk shallowly impressed before middle; occasionally there 
are one or two shallow foveate impressions on each side of disk, but they 
are never so large or so deep as in erebeci; surface as on vertex but more 
distinctly granular. Elytra obsolescently rugose, subimpunctate. Hind 
tibiae with a dense apical fringe of long setae. 

Male. Antennae (Fig. 93) reaching one or two segments beyond base 
of pronotum; segments IV-VI greatly incrassate; IV strongly expanded 
on dorsal side; V and VI each very strongly produced on dorsal side in 
the form of a heavy, blunt process which curves mesad; VII enlarged but 
only slightly distorted; VIII-X much longer than wide. Posterolateral 
surface of middle tibiae deeply impressed and densely pubescent apically 
( Fig. 192 ) . Sixth abdominal sternum as in Figure 250; emargination often 
weakly humped at middle. Genitalia as in Figure 326; ventral hooks of 
aedeagus unusually variable. 

Female. Antennae as in Figure 136, a little more strongly clavate than 
in erebea; segments IV-VII about as wide as III; IX and X at least 
slightly longer than wide. Middle tibiae moderately bowed. 

Type locality. Of eucera, Mexico or Guatemala; of spectabilis, Mexico. 

Geographic distribution. Mountains and high plateaus of Central 
America and Mexico from western Panama to Durango and central east- 
ern Coahuila. (See Fig. 37.) 

The northernmost record of this species that I can accept without 
reservation is Ciudad de Valles, San Luis Potosi. A record from Monclova, 
Coahuila, based on a female collected by E. Palmer (F. G. Werner col- 
lection), has been accepted, but as an extension of range of nearly 
400 miles is involved, verification is desirable. Published records of eucera 
from Texas (Horn, 1873; Dillon, 1952) are at best doubtful. Dillon's 
record was based on an old specimen from the C. V. Riley collection 
bearing the label "Tex." Horn recorded the species from "Texas near 
the Mexican border." 

Seasonal distribution. This species has been collected in every month 
from May to December, inclusive, with two records in February (states 
of Mexico and San Luis Potosi). 

Records. No data, 4. MEXICO: Country label only, 15. Chiapas: 
Finca La Isla (not located), 1. Coahuila'. Monclova, 1. Colima: Zapotlan 
(not located) (Champion, 1892). Distrito Federal: District label only, 1; 
Coapa, May, July, August, 5; Ciudad Madera, September, 1; Mexico 
City, 3 (1 in August); San Angel, 1; San "P." de los Pinos (not located), 
August, 1; Tacuba, 1; Tacubaya, September, 1; Valley of Mexico, August, 



146 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Llftta 




Fig. 37. Distribution of Lytta eucera. 



3. Durango: Sierra de Durango, 1. Guanajuato: State label only, 1; Silao 
(Duges, 1869). Hidalgo: Atotonilco el Grande (Duges, 1869); Pachuca, 
July, 1. Jalisco: Tlaquepaque, August, 5. Mexico: Chapingo, September, 
1; Jilotepec (Herrera and Mendoza, 1866); San Rafael, August, 2; Temes- 
caltepec, 3 (2 in February); Texcoco, August, 2. Michoacdn: Morelia, 1; 
Tacambaro [de Codallos] (Champion, 1892); Tancitaro, 6000 ft. and 6600 
ft., July, 14. Morelos: Cuernavaca (Champion, 1892); Puente de Ixtla, Sep- 
tember, 1; Tepoztlan, June, October, 2. Oaxaca: [La] Parada (Champion, 
1892); Oaxaca [de Juarez], September, 13. Puebla: Atl[ixco], October, 1; 
Cholula (Champion, 1892); Esperanza (Champion, 1892); Huauchinango 
(Herrera and Mendoza, 1866); [Izucar de] Matfamoros], November, 1; 
Puebla, May, July, 50; Techamachalco, June, August, 2. San Luis Potosi: 
C[iudad de] Valles, February, 1; Huasteca Potosina (Duges, 1889). 
Veracruz: Jalapa (Champion, 1892); Orizaba (Champion, 1892). State 
unknown: Finca Gibralter, September, 1. 

GUATEMALA: Country label only, 7; Cerro [Volcan] Zunil, 4000 ft.- 
5000 ft., 3; Chicazoa, Finca El Naraujo, 5000 ft., July, 1; Chimaltenango, 
2; San Jeronimo, 1. 

COSTA RICA: Country label only, 8. 



SYSTEMATICS AND PHYLOGENY 147 

PANAMA: Country label only, 1; Boquete, December, 3; Volcan de 
Chiriqui, 4000 ft.-6000 ft., 1. 

Remarks. From my present study it appears that the variation in this 
species is nongeographic in nature, but much additional data are required 
before this point can be decided with any degree of certainty. In a group 
where color pattern is otherwise stable intraspecifically, variation in the 
color of the head of eucera is noteworthy. Variation in the form of the 
aedeagus is considerably greater in eucera than in any of the other 
species of Lytta I have seen with the exception of quadrimacuhta. 

The first instar larva of eucera has been described by me in a separate 
publication (Selander, 1959). 

The present location of the types of eucera and spectabilis is unknown 
to me. Spectabilis has long been regarded as a synonym of eucera, and 
on the basis of its description, this synonymy seems valid. 

Bionomics. Herrera and Mendoza (1866) recorded eucera from the 
following three species of Cucurbitaceae : Sicyos angulata (as Scicyos 
angularis [sic]), Sechium edule (as Sc. edulis [sic]), and Cucurbita 
moschata (as Pepo macrocarpa). A specimen from Tacuba and one from 
Coapa, both localities in the Distrito Federal of Mexico, are from un- 
specified cucurbits. A specimen from Chapingo, state of Mexico, is re- 
corded (probably accidentally) from clover (trebol) (Leguminosae). 

Near the city of Puebla, Puebla, in 1957 I collected a series of adults 
of eucera from an unidentified species of Cucurbitaceae. I failed to find 
any specimens feeding on other species of plants in the area. Most of 
the adults were subsequently confined in an attempt to obtain eggs. 
Over a period of a week they fed solely on the cucurbit, although on sev- 
eral occasions I offered them specimens of Argemone platyceras ( Papa- 
veraceae) and several species of Compositae. Both flowers and leaves of 
the cucurbit were eaten. 

MUTILATA GROUP 

Male antennal segments IV and V or IV-VI incrassate, either ex- 
panded or produced on dorsal side. Female antennae heavier than in 
Eucera Group and less strongly clavate. Scutellum impunctate, glabrous. 
Under surface sparsely clothed with short pubescence. Setae of male 
trochanters short, erect, bristle-like ventrally. Male middle femora flat 
or concave on anterior side. Male middle tibiae enlarged apically, variously 
modified but not contorted or excavate. Male fore and middle tarsi slightly 
expanded in relation to those of female. First segment of male middle 
tarsi strongly distorted, very densely pubescent. Tarsal pads well de- 
veloped, very dense, cushion-like. Abdominal sterna as in Eucera Group. 
Male genitalia with gonostyli comparatively heavy, lobed laterally before 



148 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

apex, as in Figure 329; apex of aedeagus bluntly rounded; dorsal hook 
strongly recurved. 

Remarks. This group contains the species sanguined, erythrothorax, 
mutilata, and cardinalis. 

Key to Species 

1. Entirely orange; segments IV and V of male antennae enlarged but 

not strongly distorted ( Fig. 94 ) sanguinea ( p. 148 ) 

Color not as above; segment IV of male antennae strongly expanded, 
V very strongly, subacutely produced on dorsal side apically 2 

2. Pronotum red with a pair of large black discal spots; male fore 
femora with a notch and tuft of setae on ventral side near middle; 
segment VI of male antennae never distorted. Southern Sonora south- 
ward erythrothorax (p. 150; 

Pronotum immaculate red or black; male fore femora unmodified; 
segment VI of male antennae often distorted 3 

3. Head and pronotum red, elytra black mutilata (p. 152) 

Black with red elytra cardinalis ( p. 155 ) 

Lytta (Adicolytta) sanguinea Haag-Rutenberg 

Lytta sanguinea Haag-Rutenberg, 1880, Deutsche Ent. Zeitschr., vol. 24, 
p. 35. 

Cantharis sanguinea, Champion, 1892, Biol. Centrali-Americana, Coleoptera, 
vol. 4, pt. 2, p. 442, pi. 20, fig. 19. 

Orange. Elytra paler than head and pronotum. Antennal segments VI- 
XI (females) or VII-XI (males) black. Wings yellow. Under surface 
sometimes suffused with piceous. Pubescence pale except for some black 
setae on tarsi. Length: 16 mm.-27 mm. 

Head slightly longer than wide; sides parallel or slightly divergent 
above eyes; tempora broadly rounded; vertex tumid; surface smooth, 
moderately shiny, very sparsely punctate. Pronotum lacking definite 
lateral angles, a little longer than wide, widest at anterior third or fourth; 
disk usually broadly, distinctly impressed before middle; surface coarsely 
but not strongly alutaceous, more coarsely punctate than head. Elytra 
rugose punctate; nervules very clearly marked. 

Male. Antennae (Fig. 94) reaching base of pronotum; segments IV-VI 
moderately incrassate, weakly expanded on dorsal side, smooth on mesal 
surface, with some distortion evident in that V and VI are shallowly 
excavate for basal half dorsomedially; VII-X distinctly longer than wide. 
Modified setae of fore trochanters forming a small tuft on ventral margin 
near base. Fore femora excavate on ventral side for apical two-fifths. 
Middle femora flat or shallowly concave on anterior side, anteroventral 



SYSTEMATICS AND PHYLOGENY 



149 



edge well defined. Middle tibiae sinuate, a little enlarged apically. Pale 
pubescence (pad) of first segment of hind tarsi limited to apex. Sixth 
abdominal sternum as in Figure 251. Genitalia as in Figure 327. 

Female. Antennae as in Figure 137; segments VIII-X about one-fifth 
wider than III. Middle tibiae weakly bowed. First segment of middle and 
hind tarsi entirely black pubescent beneath. 

Type locality. Mexico. 

Geographic distribution. Valley system of the Rio Balsas and its tribu- 
taries from western Michoacan to Morelos and eastern Guerrero. (See 
Fig. 38.) 

Seasonal distribution. July 6 — September. 

Records. MEXICO: Country label only, 1 (type). Guerrero: State 
label only, 1; Iguala, September, 1; Mochitlan (Champion, 1892); Rio 
Balsas [Balsas], 9. Michoacan: Apatzingan [de la Constitution], 1200 
ft., July, August, 9. Morelos: 16 mi. south of Cuernavaca, ±1600 ft, 
July, 20; Jojutla, August, 3. 

Remarks. The immaculate orange color easily distinguishes this species 
from all other species of Lytta. It is the only species of its group in 
which antennal segment V is not strongly produced in the male. 




Fig. 38. Distribution of Lytta sanguined. 



150 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lljttd 

The type of sanguined is in the Zoologische Staatssammlung in Munich 
( examined ) . 

Bionomics. The series of specimens from 16 mi. south of Cuernavaca, 
Morelos, was collected in 1955 from two species of Convolvulaceae grow- 
ing in an area of disturbed ground at roadside. Eighteen of the specimens 
were found on an undetermined, purple-flowered species of Ipomoea. 
Feeding on this plant seemed to be confined to the corolla, mainly along 
the lip. Care was necessary in collecting the beetles as they tended to 
select deeply shaded flowers near the ground where they were easily over- 
looked. Two individuals were found on the flowers of Operculina orni- 
thopoda, but feeding was not observed. Plant identifications were made 
by F. Miranda. 

Lytta (Adicolytta) erythrothorax (Herrera and Mendoza) 

Cantharis erytrotora Herrera and Mendoza, 1867, Gaceta Med. Mexico, 
vol. 3, p. 14. 

Cantharis erytrotorax, Duges, 1869, La Naturaleza, vol. 1, p. 166 (emenda- 
tion). 

Cantharis stigmata Duges, 1881, Gaceta Med. Mexico, vol. 5, pi. 4, fig. 10. 
Not Duges, 1869, La Naturaleza, vol. 1, p. 159. 

Cantharis erythrothorax, Horn, 1885, Trans. Amer. Ent. Soc, vol. 12, p. 108 
(emendation). Duges, 1889, An. Mus. Michoacano, vol. 2, p. 102. Champion, 
1892, Biol. Centrali-Americana, Coleoptera, vol. 4, pt. 2, p. 439, pi. 20, fig. 13. 

Black. Head, except for neck and epistoma, red or orange. Pronotum 
red or orange, finely margined with black and with a large black spot 
on each side of disk at middle ( very rarely absent ) ; spots variable in size 
and shape. Scutellum and midline area of metasternum occasionally red- 
dish. Wings brown. Length: 17 mm. -23 mm. 

Head as wide as to one-tenth wider than long; sides nearly parallel 
above eyes; tempora broadly rounded; vertex tumid; frontal area pos- 
sessing a broad, oblique impression on each side extending from tip 
of midline impression to mesal margin of eye; surface smooth, shiny, 
very sparsely punctate. Pronotum generally without distinct lateral 
angles, as wide as or slightly wider than long, widest anteriorly; disk 
broadly, deeply, transversely impressed before middle, weakly impressed 
on midline basally; surface as on vertex but a little more densely punc- 
tate at sides. Elytra rugose, subimpunctate. 

Male. Antennae (Fig. 95) reaching base of pronotum; segments IV 
and V incrassate; IV strongly expanded on dorsal side, smooth on mesal 
side, two-fifths longer than wide; V very strongly, subacutely produced 
on dorsal side apically, smooth and excavate on dorsomesal side basad 
of apex of process (but not so strongly excavate as in mutilata and 
cardinalis); smooth areas of IV and V generally reddish; VI not modified; 
VI-X obviously longer than wide. Modified setae of fore trochanters 



SYSTEMATICS AND PHYLOGENY 



151 



forming a well-marked tuft on ventral margin near base. Fore femora 
with a notchlike groove and a tuft of coarse setae on ventral side near 
middle. Middle femora compressed; anterior side excavate from near base 
to apex, with an elevated area along basal margin and part of ventral 
margin which is very densely clothed with long pubescence. Middle tibiae 
enlarged apically, flat and subglabrous apically on mesal side, densely 
padded with moderately long, semierect pubescence apically on anterior 
side. Pale pubescence (pad) of first segment of hind tarsi limited to apex. 
Sixth abdominal sternum as in Figure 252; emargination sometimes 
weakly humped at middle. Genitalia as in Figure 328. 

Female. Antennae as in Figure 138; segments V-X at least slightly 
longer than wide. Middle tibiae weakly bowed. Pale pubescence (pad) 
of first segment of hind tarsi limited to apex or absent. 

Type locality. Of enjthrothorax, Taxco [de Alarcon], Guerrero; of 
stigmata, probably Tupataro, Guanajuato. 

Geographic distribution. Southern Sonora to central Jalisco and Colima; 
thence southeast to Veracruz and eastern Guerrero. ( See Fig. 39. ) 

A label notation of "Vera Cruz" for a specimen of enjthrothorax in the 
Duges collection has been interpreted as referring to the state rather 




Fig. 39. Distribution of Lijtta enjthrothorax. 



152 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

than the city of Veracruz. A specimen in the United States National 
Museum is mislabeled Texas (H. Ulke collector). 

Seasonal distribution. July 10 — October 16; also one record for March 4. 

Records. No data, 10. MEXICO: Country label only, 13. Colima: 
State label only, 1; Colima, September, 2; Tonila, 1. Guanajuato: Tupataro 
(Duges, 1889); 6 mi. northwest of Yuriria, July, 1. Guerrero: Amula, 
6000 ft., August, 1; Michapa (not located; possibly refers to Los Reyes 
Michapa, Oaxaca), September, 1; Taxco [de Alarcon], August, 1. Jalisco: 
Guadalajara, March, 2; Vulcano [Volcan de] Colima, 6. Michoacdn: 
Morelia, 1. Morelos: Cuernavaca, 8; Jalastoc [Jaloxtoc], October, 1; Te- 
poztlan, August, 5. Nayarit: Vicinity of Compostela, August, 1; San Bias, 
September, 1. Sinaloa: Mazatlan, September, 4; Pericos, August, 30; 
Venodio (not located; possibly an error for Venadillo), August, 1. Sonora: 
San Bernardo, July, 2. Veracruz: State label only, 1. 

Remarks. The only species with which erythrothorax might be con- 
fused is mutilata, which differs from it most conspicuously by the char- 
acters given in the key. The pronotal spots are absent in one specimen of 
erythrothorax examined, and Champion (1892) noted the same condition 
in one of his specimens. 

The types of both erythrothorax and stigmata have been lost. As neo- 
type of erythrothorax I have designated a male in the British Museum 
( Natural History ) from Amula, Guerrero. As neotype of stigmata I have 
designated a male in the United States National Museum labeled Mexico 
(Duges collection). 

Bionomics. The series of specimens from Pericos, Sinaloa, and the single 
specimen from near Yururia, Guanajuato, were collected on unidentified 
grasses. 

Lytta (Adicolytta) mutilata (Horn) 

Cantharis mutilata Horn, 1875, Trans. Amer. Ent. Soc, vol. 5, p. 155; 1883, 
Trans. Amer. Ent. Soc, vol. 10, p. 310. Snow, 1883, Trans. Kansas Acad. Sci., 
vol. 8, p. 43. Champion, 1892-93, Biol. Centrali-Americana, Coleoptera, vol. 
4, pt. 2, pp. 439, 463, pi. 21, fig. 21. Horn, 1894, Proc. California Acad. Sci., 
ser. 2, vol. 4, p. 356. Cockerell, 1898, New Mexico Agr. Exp. Sta. Bull. 28, 
p. 173. Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 296. Fall and Cockerell, 
1907, Trans. Amer. Ent. Soc, vol. 33, p. 210. 

Lytta mutilata, Vaurie, 1950, Amer. Mus. Novitates, no. 1477, p. 40. Dillon, 
1952, Amer. Midland Nat., vol. 48, p. 349. MacSwain, 1956, Univ. California 
Publ. Ent., vol. 12, p. 86, pi. 15. 

Black. Head, except epistoma, lower margin of frontal area, antennal 
callosities, and often a narrow area along dorsomesal margin of eyes, 
red or orange. Pronotum red or orange, generally finely margined with 
black and with a black blotch on each of the inflexed sides above coxae. 



SYSTEM ATICS AND PHYLOGENY 153 

Scutellum generally orange. Wings colorless, with pale brown veins. 
Midline area of metasternum usually reddish. Length: 12 mm. -24 mm. 

Head as wide as long; sides weakly divergent above eyes; tempora 
broadly rounded; vertex and upper frontal area smooth, shiny, very 
sparsely punctate. Pronotum with obtuse lateral angles or none, slightly 
wider than long to slightly longer than wide, widest before middle; disk 
flattened or transversely impressed before middle, frequently with a fovea 
on each side just behind middle; surface as on vertex but less finely 
micropunctate and a little less shiny. Elytra rugose, impunctate. 

Male. Antennae (Fig. 96) reaching base of pronotum; segments IV 
and V incrassate; IV strongly expanded on dorsal side, smooth on mesal 
side, one-fifth to one-third longer than wide; V very strongly, subacutely 
produced on dorsal side apically, smooth and excavate on dorsomesal 
side basad of apex of process; VI often weakly distorted, not enlarged; 
smooth areas of IV and V generally dark orange or reddish; VII-X 
nearly as wide as long. Fore trochanters with ventral margin strongly 
angulate basally; modified setae forming a definite row on ventral margin. 
Fore femora unmodified. Middle femora compressed; anterior side ex- 
cavate for apical two-thirds, with an elevated area basally which is very 
densely clothed with long pubescence (area less extensive than in erythro- 
thorax). Middle tibiae enlarged apically, nearly straight, flat and glabrous 
apically on mesal side, densely padded apically on anterior side with 
moderately long, semierect pubescence. Pale pubescence (pad) of first 
segment of hind tarsi limited to apex or absent. Sixth abdominal sternum 
as in Figure 253, or with a definite apex indicated at middle of emargina- 
tion. Genitalia as in Figure 329. 

Female. Antennae as in cardinalis (see Fig. 139) but segments V-X 
progressively slightly wider and VI-X as wide as long. Middle tibiae 
strongly bowed. Pale pubescence (pad) absent on first segment of hind 
tarsi. 

Type locality. Arizona. 

Geographic distribution. Northeastern Durango and central eastern 
Coahuila to southern Arizona and southern New Mexico, extending west 
to the Pacific Coast of California near the Mexican border and thence 
south to southern Baja California Norte. (See Fig. 40). 

Seasonal distribution. June 28 — October 5. 

Records. UNITED STATES: Arizona: Douglas, August, 4; Marana, 
August, 1; Nogales, August, 1; Patagonia, August, 1; Phoenix, 10 (7 in 
September ) ; Postvale, August, 2; Redrock, September, 2; Thatcher, Aug- 
ust, 84; Tumacacori Mission, 3200 ft., August, 8; Tucson, July-October, 
22; Willcox, August, 1. California: La Jolla, 1. New Mexico: State label 



154 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ll/tta 




Fig. 40. Distribution of Lytta mutilata. 



only, 2; 10 mi. north of Columbus, 4100 ft., September, 1; Deming, Sep- 
tember, 1; Mesilla (Fall and Cockerell, 1907); Mesilla Park, August, 1; 
Socorro (Snow, 1883). Texas: State label only, 4; Alpine, June, August, 
10; El Paso, 4000 ft., September, 1; Pine Springs, August, 47; Presidio, 
August, 44. 

MEXICO: Baja California Norte: San Julio (Horn, 1894). Chihuahua: 
9 mi. north of Camargo, September, 1; Salaices, 5200 ft., September, 
October, 2; Kilometer 36, [between] Santa Barbara [and] Ojito [in 
Durango], 6900 ft., August, 17. Coahuila: Monclova (Champion, 1892). 
Durango: Villa [Ciudad] Lerdo (Champion, 1892). 

Remarks. Lytta mutilata is the only representative of Adicolytta de- 
finitely known to occur in the United States. Although apparently most 
closely related to cardinalis, it more nearly resembles peninstdaris and 
erythrothorax in color. From peninsularis it is readily separated by the 
characters of the Mutilata Group and from erythrothorax by the char- 
acters given in the key to the species of the Mutilata Group. 

The first instar larva of mutilata has been described by MacSwain 



SYSTEM ATICS AND PHYLOGENY 155 

( 1956 ) . The type of the species is in the collection of the Academy of 
Natural Sciences of Philadelphia. 

Bionomics. Specimens have been collected on Cuscuta (Convolvula- 
ceae) at Pine Springs, Texas, by J. W. MacSwain and at Redrock, Pinal 
County, Arizona, by F. F. Bibby and J. W. Warren. 

Lytta (Adicolytta) cardinalis Chevrolat 

Lytta cardinalis Chevrolat, 1834, Coleopteres du Mexique, fasc. 1, p. [25]. 

Cantharis cardinalis, Chevrolat, 1834, Coleopteres du Mexique, fasc. 3, p. 
[19]. Duges, 1869, La Naturaleza, vol. 1, p. Ill, pi. 1, fig. 12. Horn, 1885, 
Trans. Amer. Ent. Soc., vol. 12, p. 111. Duges, 1889, An. Mus. Michoacano, 
vol. 2, p. 102. Champion, 1892, Biol. Centrali-Americana, Coleoptera, vol. 4, 
pt. 2, p. 438, pi. 20, fig. 12. 

Black. Head with a diamond-shaped orange frontal spot. Elytra a rich 
dark red. Wings yellow. Length: 13 mm. -25 mm. 

Head barely longer than wide; sides parallel above eyes; tempora 
broadly rounded; dorsal margin weakly rounded; surface smooth, moder- 
ately shiny, very sparsely punctate; center of frontal spot elevated. Pro- 
notum with obtuse lateral angles, as wide as long, widest just before 
middle; disk transversely impressed before middle, moderately shiny, 
finely, weakly, irregularly alutaceous, center before middle subimpunc- 
tate, glabrous. Elytra rugose, subimpunctate. 

Male. Antennae (Fig. 97) nearly reaching base of pronotum; segments 
IV and V incrassate; IV strongly expanded on dorsal side, smooth on 
mesal side, about one-third longer than wide; V very strongly, subacutely 
produced on dorsal side apically, smooth and excavate on dorsomesal 
side basad of apex of process; VI often weakly distorted, not enlarged; 
VII-X distinctly longer than wide. Fore trochanters not strongly angulate 
on ventral margin; modified setae forming a definite row on ventral mar- 
gin, setae increasing in length toward base of trochanter. Fore femora un- 
modified. Middle legs modified as in mutilata. Pale pubescence (pad) of 
first segment of hind tarsi limited to apex. Sixth abdominal sternum 
shallowly emarginate. Genitalia as in Figure 330. 

Female. Antennae as in Figure 139; segments V-X not becoming per- 
ceptibly wider, each longer than wide. Middle tibiae strongly bowed. 
Pale pubescence (pad) absent on first segment of hind tarsi. Sixth ab- 
dominal sternum as in Figure 281. 

Type locality. Tehuacan, Puebla. 

Geographic distribution. Southern Puebla, northwestern Oaxaca, and 
eastern Guerrero to central Jalisco, extending east into Veracruz, prob- 
ably in the region adjacent to southern Puebla. (See Fig. 41.) 

Extension of the range of this species to Jalisco is based on a specimen 
in the Duges collection. Labels of "Vera Cruz" for three additional 



156 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 




Fig. 41. Distribution of Lytta cardinalis. 



specimens in the Duges collection have been interpreted as referring to 
the state rather than the city of that name. 

Seasonal distribution. September 16 — November. 

Records. No data, 2. MEXICO: Country label only, 2. Guerrero: Mo- 
chitlan (Champion, 1892). Jalisco: Guadalajara, 1. Morelos: Alpuyeca, 
1100 m., October, 9; Cuautla [Morelos], 1; Xochicalco (not located), 
October, 1; Yautepec, 9. Oaxaca: Huajuapan de Leon, September, 4; 
Tamazulapan, September, 1. Puebla: Chiantla [error for Chiautla], 1; 
Matamoros Iziicar [Iziicar de Matamoros], 9 (5 in November); Puebla, 1; 
Techamachalco, October, 2; Tehuacan (Chevrolat, 1834). Veracruz: State 
label only, 3. 

Remarks. This attractive species is easily distinguished from all other 
New World species of Lytta by its distinctive color pattern. Morpho- 
logically it is most like mutilata. 

The location of the type of cardinalis is unknown to me. 

Bionomics. According to Chevrolat ( 1834), adults were found in Puebla 
on "une plante grimpante." Dr. C. Bolivar y Pieltain has informed me 
that he has seen adults on a species of white-flowered Convolvulaceae in 
Morelos. 



SYSTEM ATICS AND PHYLOGENY 157 

Subgenus POREOSPASTA Horn, new status 

Poreospasta Horn, 1868, Trans. Amer. Ent. Soc, vol. 2, p. 139. MacSwain, 
1943, Ann. Ent. Soc. America, vol. 36, p. 360. 

Small to moderately large beetles. Colors black or metallic (often bril- 
liant) green, blue, or purple; head and/or pronotum sometimes partially 
or entirely orange. Male antennae usually long, longer than in female, 
reaching or extending beyond base of pronotum (except in sublaevis); 
intermediate segments incrassate or not, never excavate or produced; if 
intermediate segments incrassate, male hind trochanters spined or angu- 
late, or gonostyli of male genitalia hairy, or both. Female antennae 
clavate. Elytra rugose or (sublaevis) smooth. Male fore tibiae each with 
two spurs, or if posterior spur absent (nigrocyanea and some moerens), 
then hind trochanters spined. Pygidium not produced. Male genitalia 
with each gonostylus bearing a mesal ( or mesoventral ) hook, or if hooks 
absent, gonostyli clothed ventrally with long setae; aedeagus with one 
or two ventral hooks (both absent in childi). 

Type species. Nomaspis sublaevis Horn, by subjective synonymy with 
Poreospasta polita Horn (preoccupied); fixed by monotypy. 

Remarks. In North America this subgenus contains 21 species, divided 
into two groups. Its range includes most of the western United States 
and adjacent areas in southwestern Canada and northwestern Mexico 
(Figs. 7-8). In addition to the North American representatives, there are 
at least four Palearctic species (luteovittata Kraatz, clematidis Pallas, 
poeciloptera Semenov, and fairmairei Borchmann) that fall within the 
limits of the subgenus as presently defined. Phylogenetically Poresospasta 
seems to be nearer to the subgenera Lytta and Pomphopoea than to 
either Paralytta or Adicolytta. 

Adults of Poreospasta feed on a wide variety of herbaceous plants. 
Larval hosts have been recorded for three species ( Table I ) . 

Five groups of Poreospasta may be recognized: the Luteovittata, Fair- 
mairei, and Clematidis groups in the Palearctic region and the Stygica 
and Moerens groups in the Nearctic region. 

The Luteovittata Group (luteovittata) may be defined as follows: 
Frontal spot present. Head subtriangular. Antennae moniliform clavate 
in both sexes. Male legs unmodified; all tibiae with two spurs each. Male 
sixth abdominal sternum with a lightly sclerotized area. Male genitalia 
with each gonostylus bearing a rather poorly developed hook meso- 
ventrally at apex, clothed ventrally with long setae; aedeagus with a 
single ventral hook. Luteovittata closely approaches the species moerens 
in most characters. 

The Fairmairei Group (fairmairei) has the following combination of 



158 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytttt 

characters: Frontal spot present. Head subquadrate. Antennae filiform; 
segments closely articulated; male intermediate segments not incrassate. 
Male legs unmodified; all tibiae with two spurs each; hind tibial spurs 
both slender, flattened, sticklike. Male sixth abdominal sternum without 
a lightly sclerotized area. Male genitalia with each gonostylus bearing 
a mesal hook, clothed ventrally with long setae; aedeagus without ven- 
tral hooks. 

The Clematidis Group (clematidis and poeciloptera) has these char- 
acters: Frontal spot absent. Head quadrate. Antennae moniliform; male 
intermediate segments not incrassate. Male first fore tarsal segment 
modified ( poeciloptera ) or not; legs otherwise unmodified; all tibiae with 
two spurs each. Male sixth abdominal sternum without a lightly sclero- 
tized area. Male genitalia with gonostyli each bearing a mesal hook, 
clothed ventrally with long setae; aedeagus with two ventral hooks, the 
subapical one small. 

The two North American groups of Poreospasta I have recognized were 
recently delimited by MacSwain (1956) in his classification of the first 
instar larvae of the Meloidae. According to MacSwain, first instar larvae 
of the Moerens and Stygica ( = Polita ) groups are separable on the basis 
of characters of the mandibles (distinct teeth present in the Moerens 
Group) and maxillae (third segment with lateral margins usually con- 
verging in the Moerens Group, never so in the Stygica Group). Mac- 
Swain's diagnosis of the adults of the Moerens Group is inaccurate, it 
should be noted, in that it fails to recognize the presence of mesal hooks 
on the male gonostyli of some of the included species. 

Of the two North American groups, the Stygica Group is taxonomically 
the more difficult and at the same time the more interesting. In the 
course of the present study I have spent a disproportionately large 
amount of time in the analysis of this group, without arriving at wholly 
satisfactory results as to species limits or species relationships. The group 
is confined largely to California and is remarkable for the high degree 
of sympatry shown by its species. In both the larval stage and the adult 
stage the species exhibit a general uniformity of structure that makes it 
impossible to arrange them in a phylogenetic system with any great 
degree of confidence. In short, the group has undergone considerable 
speciation with relatively little differentiation. MacSwain, who also re- 
marked at the difficulty of determining relationships among the species, 
has suggested some groupings on the base of larval morphology, but I 
fail to find support for these in characters of the adults. The relationships 
suggested in Figure 42 are static ones based entirely on adult morphology 
and are to be regarded as highly tentative. The morphological basis for 
the arrangement is largely outlined in the key to species (below). 

The species rathvoni, chloris, comans, and stygica form a complex 



SYSTEMATICS AND PHYLOGENY 



159 



that apparently cannot be understood without intensive field studies. 
The present treatment of this complex is therefore provisional, serving 
mainly to outline some of the evolutionary and taxonomic problems 
awaiting study. 

The Moerens Group, which constitutes the second North American line 
of Poreospasta, seems to have undergone much of its early evolution on 
the Pacific Coast, where its more primitive species are now found. In the 
course of its development it has spread eastward and northward into 
the Rocky Mountains. Adults of the group are specialized with respect 
to other members of the subgenus in having the male hind trochanters 
generally angulate or spined, the male first fore tarsal segment distinctly 
cultriform, the male fifth abdominal sternum deeply emarginate, and the 
male sixth abdominal sternum with a lightly sclerotized median area. In 
six of the nine species of the group the intermediate segments of the 
male antennae are swollen to the extent that they are in part as wide as 
or wider than the distal segments. This character suggests the possibility 
of a relationship with the subgenus Paralytta, but the weight of evidence 
favors the conclusion that the similarity between the two is convergent. 
Whether the incrassate condition of the antennae is a primitive or special- 
ized character within the Moerens Group itself is questionable. If it is 
primitive, then according to my interpretation of the phylogeny (Fig. 
42), moerens and navajo on the one hand and morosa on the other must 



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Fig. 42. Phylogeny of the North American species of Poreospasta. 



160 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

have lost it independently. Conversely, if it is a specialized character, 
it presumably arose independently on two or even three occasions. 

First instar larvae has been described by MacSwain (1956) for one of 
the four species of the Moerens Subgroup, two of the three species 
of the Cyanipennis Subgroup, and both species of the Childi Subgroup. 
Evidence from the larval morphology fully supports the division of the 
group into three subgroups suggested by adult morphology. 

The Moerens Subgroup contains all those species of the group which 
retain the mesal hooks on the male gonostyli. It seems likely that the 
most primitive of these is moerens. The species navajo is similar to 
moerens in most characters and is apparently a derivative of it. Insperata 
differs from moerens and navajo mainly in its incrassate male antennae, 
distinctly spined male hind trochanters, and elongate male sixth ab- 
dominal sternum and genitalia. With respect to the first two characters it 
is like nigrocyanea, but both characters are very likely primitive for the 
subgroup, and I feel more inclined to regard insperata as a separate 
offshoot of the original ancestor of the subgroup than as a close relative 
of nigrocyanea. 

The species nigrocyanea is itself a "missing link" connecting moerens 
and navajo with the species of the Cyanipennis Subgroup. It resembles 
moerens and navajo in having the male first fore tarsal segment unmodi- 
fied and in retaining mesal hooks on the male gonostyli, while at the 
same time it has the general facies of the Cyanipennis Subgroup and 
almost precisely the same incrassate antennal type. It is not, however, a 
direct ancestor of the Cyanipennis Subgroup as the male is more special- 
ized than any species of that subgroup in having only a single fore tibial 
spur. 

The Cyanipennis Subgroup includes three similar, bright metallic 
species in which the mesal hooks of the male gonostyli have been lost 
and in which the first segment of the male fore tarsi is strongly distorted. 
Within the subgroup the species may be arranged on the basis of static 
relationships in the order shown in Figure 42. 

In the present state of our knowledge, the Childi Subgroup is most 
logically regarded as a separate derivative of the ancestor of the Moerens 
Group. It resembles the Cyanipennis Subgroup in characters of the male 
genitalia (gonostyli lacking mesal hooks and clothed with long setae). 
However, there is little else to suggest a special relationship between 
the two subgroups, and I regard the genitalic similarity as probably 
convergent. 

Key to North American Groups 

1. Male intermediate antennal segments not incrassate, never wider 
than distal segments; male hind trochanters unmodified; male first 



SYSTEMATICS AND PHYLOGENY 161 

tarsal segment not cultriform; male fifth abdominal sternum not deeply 
emarginate; male sixth sternum not modified as below; gonostyli each 

bearing a mesal hook; aedeagus with one or two ventral hooks 

Stygica Group ( p. 161 ) 

Male intermediate antennal segments incrassate or not; male hind 
trochanters usually spined or angulate on ventral margin (always 
so if antennae not incrassate); male first tarsal segment cultriform; 
male fifth abdominal sternum deeply emarginate; male sixth sternum 
elongate, very deeply, narrowly, triangularly emarginate, with a lightly 
sclerotized area medianly, clothed with long setae; gonostyli with or 

without mesal hooks; aedeagus with one ventral hook or none 

Moerens Group ( p. 193 ) 

STYGICA GROUP 

Head usually with a pale (orange) frontal spot. Head sparsely micro- 
punctate, finely to moderately coarsely, usually rather sparsely punctate. 
Male antennae slender, except in sublaevis reaching at least base of pro- 
notum; segments much more elongate than in female; intermediate seg- 
ments relatively wider than in female but not wider than distal segments 
and not appearing incrassate. Legs slender, unmodified. Male hind tro- 
chanters neither spined nor angulate on ventral margin. Male middle 
tibial spurs both spiniform. Outer hind tibial spurs usually thickened, at 
least slightly, obliquely or very obliquely truncate; inner spur spiniform, 
flattened or very obliquely truncate. Male fore and middle tarsi very 
slightly heavier than in female. Tarsal claws slender. Male fifth abdominal 
sternum not deeply emarginate. Male sixth sternum not elongate, lacking 
a median lightly sclerotized, long-pubescent area, not clothed with long 
pubescence on lateral lobes; emargination triangular or shallowly obtuse. 
Male genitalia with each gonostylus bearing a mesal hook, glabrous or 
essentially so; aedeagus with one or two ventral hooks, the apical one 
small when present. 

Remarks. This group contains the species sublaevis, auriculata, nigri- 
pilis, hoppingi, lugens, aeneipennis, refulgens, crotchi, chloris, rathvoni, 
stygica, and comans. It is largely confined to California, with only two 
species (auriculata and stygica) ranging outside the limits of the state 
( Fig. 7 ) . All 12 species included in it are very similar to one another in 
both adult and larval characters, and on this account no formal division 
of the group above the species level has been made. 

The adult beetles are found most commonly on flowers of Eschscholtzia 
( Papaveraceae ) and Ceanothus ( Rhamnaceae ) , although a great variety 
of other plants also serve as hosts, including particularly species of 
Leguminosae and Compositae. The principal, if not the only, food 
material utilized by the adults is pollen. 



162 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Key to Species 

1. Wingless, with shortened elytra sublaevis (p. 163) 

Wings and elytra fully developed 2 

2. Black, with dark metallic blue or green elytra; conspicously clothed 
throughout with rather long, more or less erect pubescence; male and 
female sixth abdominal sterna as in Figures 255 and 282, respectively; 
male genitalia with ventral hooks of aedeagus rudimentary (Figs. 
332c and 333c) and mesal hooks of gonostyli located nearer to mid- 
dle than to apex 3 

Not as above 4 

3. Pubescence colorless apically or entirely colorless, at least on under 
surface of body; vertex often marked with orange; male antennae 
(Fig. 255) nearly filiform, with more cylindrical segments; male 

genitalia as in Figure 332 auriculata (p. 165) 

Pubescence piceous; vertex entirely black; male antennae (Fig. 100) 
more clavate, with more moniliform segments; male genitalia as 
in Figure 333. Sierra Nevada, and coastal California north of San 
Francisco Bay nigripilis ( p. 169 ) 

4. Pronotum distinctly elongate, one-tenth to nearly one-third longer 
than wide; slender, black beetles, without metallic luster; pronotum 
with sides orange or not; outer hind tibial spur not or only slightly 

wider than inner spur 5 

Not as above 6 

5. Pronotum orange with a wide black median stripe . hoppingi (p. 171) 
Pronotum entirely black higens (p. 172 ) 

6. Pronotum orange, at least in part 7 

Pronotum black, green, blue, or metallic 9 

7. Vertex orange aeneipennis (p. 174) 

Head green except for usual small pronotal spot 8 

8. Pronotum with a large green median spot; elytra with long pubes- 
cence confined to base refulgens (p. 175) 

Pronotum entirely orange; elytra clothed throughout with long, erect 
pubescence crotchi ( p. 177 ) 

9. Elytra violet or green with a violet ( not brassy ) luster; rest of body 
green or brass-colored; pubescence colorless; head and pronotum 
distinctly microreticulate; male sixth abdominal sternum medianly 
impressed, moderately deeply, triangularly emarginate; female sixth 
sternum emarginate. Sacramento and Calaveras counties, California, 

south to Tulare County, California rathvoni (p 178) 

Not as above 10 



SYSTEMATICS AND PHYLOGENY 163 

10. Head and pronotum distinctly microreticulate, more or less satiny in 
texture; male sixth abdominal sternum (Fig. 262) relatively flat, 
shallowly, evenly emarginate; male antennal segments elongate, more 
cylindrical (Fig. 106); brilliant brassy green; pubescence colorless 
(Kern and Tulare counties, California) or piceous (central western 

California ) chloris ( p. 179 ) 

Head and pronotum smooth or relatively so; male sixth abdominal 
sternum medianly impressed, moderately deeply, triangularly emar- 
ginate; male antennal segments more moniliform; pubescence piceous 
11 

11. Hairy, conspicuously clothed throughout with long, straight, erect 

pubescence; green. Central western California comans (p. 192) 

Pubescence shorter, less erect, especially on upper surface; elytral 
pubescence very short, subrecumbent, and inconspicuous, or if 
longer, erect setae present these confined to immediate base; green, 
blue, or black stygica (p. 185) 

Lijtta (Poreospasta) subloevis (Horn) 

Nomaspis sublaevis Horn, 1868, Trans. Amer. Ent. Soc, vol. 2, p. 140. 

Poreospasta polita Horn, 1868, Trans. Amer. Ent. Soc, vol. 2, p. 139. Van 
Dyke, 1928, Univ. California Publ. Ent., vol. 4, p. 415, pi. 15, fig. 4. MacSwain, 
1943, Ann. Ent. Soc. America, vol. 36, p. 361, fig. 2. Edwards, 1951, Pan- 
Pacific Ent., vol. 27, p. 44. Not Lytta polita Say, 1824, Jour. Acad. Nat. Sci. 
Philadelphia, vol. 3, p. 302. New sxjnomjmy. 

Poreospasta sublaevis, Van Dyke, 1928, Univ. California Publ. Ent., vol. 4, 
p. 416, pi. 15, fig. 3. 

Lytta polita, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 95. 

Black. Head with a large orange spot at center above eyes, occasionally 
with occiput orange also. Pronotum often orange at sides, rarely with 
disk orange except for a pair of black spots. Length: 11 mm. -20 mm. 

Wingless, with elytra shortened and abdomen somewhat swollen. Head 
with sides straight, generally divergent; tempora well marked; dorsal 
margin weakly curved; surface finely granular to smooth, finely, sparsely, 
deeply punctate; pubescence short, scattered. Eyes very small, oval. Pro- 
notum subhexagonal, wider than long; disk even, flattened, sometimes 
with a fovea on each side at middle; surface glabrous, impunctate or very 
sparsely punctate at center of disk. Scutellum large, broad, bluntly 
rounded or truncate, generally foveate. Elytra shortened, somewhat in- 
flated, covering mesothoracic epimera at sides, exposing last two ab- 
dominal terga when abdomen is not badly shrunken; sutural margins 
touching or overlapping for basal two-thirds to three-fourths, then 
strongly divergent; surface smooth (not rugose), very finely punctate, 
subglabrous. Hind wings entirely absent. Mesothorax shortened, the 



164 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

middle coxae almost reaching metasternum, not overlapping hind coxae. 
Middle tibiae bowed. Outer hind tibial spur twice as wide as inner spur, 
expanded apically, very obliquely truncate; inner spur very obliquely 
truncate. Tarsal pads moderately well developed, undivided; first seg- 
ment of middle tarsi and all segments of hind tarsi entirely dark pubes- 
cent. Abdomen somewhat swollen; terga and sterna fully developed, 
normal. 

Male. Antennae (Fig. 98) short, reaching about middle of pronotum, 
segment III fully twice as long as wide; IV-X shorter, subequal in length, 
becoming progressively more moniliform and slightly wider. Fifth ab- 
dominal sternum feebly emarginate. Emargination of sixth sternum ( Fig. 
254) moderately deep, triangular; a shallow median sulcus from apex of 
emargination. Genitalia as in Figure 331; apical ventral hook of aedeagus 
obsolescent. 

Female. Tending to be larger than male and with more strongly swollen 
abdomen. Antennae shorter (Fig. 140). Sixth abdominal sternum shal- 
lowly, triangularly emarginate. 

Type locality. Of polita, "Southern Coast Range of California" (Horn, 
1868); of sublaevis, Fort Tejon, California. 

Geographic distribution. Western side of the Central Valley of Cali- 
fornia and adjacent hills from Alameda and San Joaquin counties south; 
thence east along the Tehachapi Mountains and north along the eastern 
edge of the Central Valley to Tulare County. ( See Fig. 45. ) 

Seasonal distribution. February 21 — July 25. 

Records. UNITED STATES: California: Alamont Pass, Alameda 
County, April, 1; Arvin, February-April, 48; 20 mi. east of Bakersfield, 
April, 7; 10 mi. north of Blackswells Corner, April, 4; Coalinga and 10 mi. 
east of Coalinga, March, April, 3; Corral Hollow, San Joaquin County, 
March, 2; Cottonwood Creek, Kern County (ambiguous; not mapped), 
March, April, 5; 5 mi. south of Democrat Springs, Kern County, April, 3; 
Ducor, March, 4; Fort Tejon (Horn, 1868); Gorman (Van Dyke, 1928); 
Hospital Canyon, San Joaquin County, March, April, 6; Ingram Canyon, 
8 mi. west of Westley, April, 3; Kettleman Hills, March, 1; Kings County, 
June, 7; Livermore, March, 4; Llanada, February, 41; Los Banos, Merced 
County, 2; Lost Hills, Kern County, March, April, 5; 30 mi. southwest 
of Mendota, April, 1; Vz mi. north of Midway, April, 38; Oro Lona, Feb- 
ruary, 2; Panoche and Panoche Creek, Fresno County, March, April, 2; 
Porterville, July, 1; Posey Creek (Van Dyke, 1928); Simmler, March, April, 
29; Tipton, March, 8; Tracy and vicinity, March, 27; Tulare Lake, March, 
19; Visalia, March, June, 3; Warthan Canyon, Fresno County, March, 4; 
Woody, March, 1. 



SYSTEMATICS AND PHYLOGENY 165 

Remarks. This is the only wingless species of Lytta. Superficially, the 
adults are very similar to those of the genus Meloe, but they are easily 
distinguished by the presence of the pale frontal spot on the head, as 
well as by the presence of mesal hooks on the gonostyli of the male 
genitalia. 

Despite extensive modification of the body form accompanying the 
loss of wings, there is no question but that sublaevis belongs in the 
present group. Credit for first recognizing its true relationships belongs 
to MacSwain (1956). 

In most specimens from the western side of the Central Valley the 
head and pronotum are quite finely sculptured, the disk of the pronotum 
is relatively flat, and the sides of the pronotum are distinctly angulate. 
The head is black except for the pale frontal spot. There is some orange 
on the pronotum in approximately half the specimens examined; gener- 
ally, this color is limited to the sides, but in a few specimens it covers 
nearly all the pronotum. Color variation does not appear to follow any 
regular geographic pattern within the limits of this western race. Along 
the southern and eastern edge of the Central Valley sublaevis is repre- 
sented by a race which differs from the western form in having the head 
and pronotum more coarsely sculptured, the disk of the pronotum more 
convex, the sides of the pronotum less strongly or not at all angulate, and 
the pronotum consistently entirely black. In two of the specimens ( Ducor 
and Visalia) the top of the vertex (as well as the usual frontal spot) is 
orange. 

Typical representatives of the two races are shown in Van Dyke's work 
(1928, pi. 15, figs. 3-4). Approximately 90 per cent of the specimens 
examined can be sorted to race with ease. However, variation in both 
races, and especially in the western one, makes a consistent separation 
of specimens impossible. Interestingly, within the range of the western 
race specimens structurally resembling the southeastern race occur more 
commonly in the north than in the south. 

The larva of sublaevis has been described (as polita) by MacSwain 
(1943 and 1956). The types of polita and sublaevis are in the collection 
of the Academy of Natural Sciences of Philadelphia. 

Bionomics. There are two records of sublaevis feeding on grass, two 
from Eschscholtzia (Papaveraceae), one from Amsinckia (Boraginaceae), 
and one from Compositae. 

Lytta (Poreospasta) auriculata Horn 

Lytta auriculata Horn, 1870, Trans. Amer. Ent. Soc, vol. 3, p. 91. Van 
Dyke, 1929, Bull. Brooklyn Ent. Soc, vol. 24, p. 131. Edwards, 1951, Pan- 
Pacific Ent., vol. 27, p. 44. MacSwain, 1956, Univ. California Publ. Ent., vol. 
12, p. 90. 



166 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Cantharis auriculata, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 113; 
1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Occas. Papers Cali- 
fornia Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc., vol. 27, p. 299. 

Black, sometimes with a faint metallic blue or green luster. Head with 
a small orange frontal spot; vertex typically orange with a black stripe 
on midline but varying from entirely orange to entirely black. Elytra 
generally metallic green or bluish green, varying to nearly black. Wings 
brown, the anal region pale. Pubescence entirely colorless or colorless 
with base piceous, that on upper surface of body varying very rarely to 
entirely piceous; pubescence erect, producing a hairy appearance. 
Length: 6 mm.-19 mm. 

Head quadrate; tempora not prominent; dorsal margin truncate; vertex 
smooth, shiny, rather finely, deeply, moderately densely to sparsely punc- 
tate; punctures separated by an average distance greater than diameter 
of a single puncture; frontal area more densely punctate and usually a 
little alutaceous; pubescence long, fine, erect, conspicuous, varying in 
different specimens from half to fully as long as fourth segment of middle 
tarsus. Pronotum (Fig. 185) as wide as or slightly wider than long, not 
transverse, generally subhexagonal, with obtuse lateral angles, but vary- 
ing to nearly circular; surface before middle like that of vertex, behind 
middle more densely punctate and sometimes weakly alutaceous; pubes- 
cence as on vertex; basal margin not strongly reflexed. Scutellum rounded. 
Elytra rugose, impunctate, sparsely, regularly clothed with semierect 
setae which are usually a little shorter than those on head. Outer hind 
tibial spur two to three times as wide as inner spur, rather acute at apex; 
inner spur obtuse. Tarsal pads well developed, undivided on fore and 
middle tarsi and on first segment of hind tarsi, tending to be divided on 
midline on second to fourth segments of hind tarsi; pale pubescence 
(pad) of first segment of hind tarsi limited to apical half to two-thirds. 
Tarsal claws (Fig. 201) heavier and more strongly curved than in re- 
lated species except nigripilis. 

Male. Antennae (Fig. 99) extending two segments beyond base of 
pronotum; distal segments at most one and one-fourth longer than wide, 
only slightly wider than intermediate segments; XI about as wide as X, 
shorter than IX and X combined. Fifth abdominal sternum shallowly, 
broadly emarginate. Emargination of sixth sternum (Fig. 255) deep, tri- 
angular; sternum lacking fine median groove, impressed between lateral 
lobes, which are rather densely pubescent. Genitalia as in Figure 332; 
gonoforceps more elongate than in nigripilis; gonostyli slender, the mesal 
hook on each located distad of midpoint between base and apex; dorsal 
margin of aedeagus abruptly emarginate near apex, which is slender; 
ventral aedeagal hooks rudimentary, the apical hook often obsolete; dorsal 
hook strongly recurved. 



SYSTEMATICS AND PHYLOGENY 



167 



Female. Antennae (Fig. 141) not reaching base of pronotum. Tarsal 
pads smaller and often in part divided; pale pubescence (pad) of first 
segment of middle tarsi limited to apex. Sixth abdominal sternum not 
impressed, either emarginate (Fig. 282) or with apex entire. 

Type locality. Kern River region, California. 

Geographic distribution. Southern California and the Sonoran Desert 
of Arizona, extending north along the Colorado River system to southern 
Utah and south onto the peninsula of Baja California. ( See Fig. 43. ) 

Seasonal distribution. March — July. 

Records. UNITED STATES: Arizona: Cortaro, March, 5; Florence 
Junction, April, 1; Fredonia, June, 5; Gila Bend, April, 2; Oatman, April, 
1; Oracle Junction, April, 1; Peach Springs, June, 1; Phoenix, March-June, 
7; Pichacho, April, 1; Sabino Canyon, March, I; Sentinel, March, 19; 
Tucson, March-May, 19; Willton, April, 1. California: Andreas Canyon, 
April, 3; Argus Mountains, May, 10; Arvin, April, 9; Baker, March, 11; 
Banning, March-June, 12; Barrett Springs, April, 6; 10 mi. north of Black- 
wells Corner, Kern County, April, 10; Borrego [Springs], March, April, 
28; Buckmans Springs, San Diego County, May, 10; Cajon Pass, June, 7; 
Camarillo, March, April, 12; Carmel, May, 44; Chilao, June, 8; Chino 




Fig. 43 (left). Distribution of Lytta auriculata. 
Fig. 44 ( right ) . Distribution of Lytta nigripilis. 



168 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Canyon, March, 1; Chocolate Mountains, Imperial County, April, 3; 
Claremont, 4; Clovis, May, 3; Coachella Valley, 1; Coalinga (and War- 
than Canyon), March-May, 13; Cronise Lake, April, 2; Crystal Lake, 
Los Angeles County, June, 1; Cuyama Valley, Kern County, May, 1; 
Daggett, April, 3; Darwin, May, 21; Deep Creek, San Bernardino 
County, May, 2; Doble, June, 1; Edison, April, 2; Elizabeth Lake, Los 
Angeles County, June, 1; El Mirador, April, May, 12; El Paso Mountains, 
April, 1; Frazier Park, May, June, 65; Frenchmans Flat, Los Angeles 
County, April, 1; Fresno, June, 7; Gavilan, April, May, 3; Gilman Hot 
Springs, May, June, 5; south of Gorgonia Pass, May, 1; Griffith Park, 
April, 6; Indio, March, 1; 5 mi. west of Indio, March, 1; 20 mi. east of 
Indio, April, 13; Joshua Tree National Monument, April, 3; Kane Springs, 
March, 1; Keeler, May, 8; Kings County, April, 5; Lancaster, April, May, 
8; Lanes Bridge, Fresno County, April, 27; Lebec, 4000 ft., May, June, 12; 
Little Lake, Inyo County, May, 2; Lone Pine, May, June, 2; Los Angeles, 
April, 3; Lost Hills and 6 mi. south of Lost Hills, April, 14; Lytle Creek, 
San Bernardino County, May, 1; Maricopa Pass, July, 2; Mendota and 
30 mi. southwest of Mendota, April, 4; Milham City (MacSwain, 1956); 
Mint Canyon, April, 12; Mojave, April-June, 7; Morongo Valley, May, 1; 
Morro Bay, San Luis Obispo County, June, 1; Mount Pinos, Kern County, 
June, 5; The Narrows, San Diego County, March, April, 3; Neenach, 
May, 1; Needles, March, 4; Niland, April, 1; Owens Valley, May, 3; 
Palm Canyon, April, 3; Palmdale, May, 1; Palm Springs, March, April, 
61; Panamint Mountains, May, 13; Panoche Hills and Panoche Valley, 
April, May, 14; Paraiso Springs, May, June, 3; Pasadena, 1; 6 mi. east of 
Pearblossom, May, 31; Pine Mountains, San Luis Obispo County, April, 
1; Pinnacles, May, 10; Poway, 2; Priest Valley, Monterey County, May, 1; 
Ramona, April, 1; Randsburg, May, 1; Riverside, April, 4; Saboba Springs, 
Riverside County, June, 1; San Bernardino, 1; San Carlos Canyon, May, 1; 
San Diego, April, May, 10; San Jacinto Mountains (Cranston Ranger 
Station, Herkey Creek, Idyllwild, 8 mi. -9 mi. west of Keen Camp, Pinon 
Flat, Ribbonwood, San Jacinto River, 3000 ft., Vandevanter Flat), May, 
June, 55; San Lucas, April, 4; San Marcus Pass, June, 2; Santa Cruz, June, 
2; Santa Monica, April, 3; Santa Rosa Mountains, 5800 ft. and 6200 ft.. 
May, June, 15; 2 mi. north of Shavers Well, Riverside County, April, 3; 
Tanbark Flat and 4 mi. west of Tanbark Flat, Los Angeles County, June, 
7; Tehachapi, May, 14; Tejon Pass, May, 2; Tulare Lake, March, 4; Ven- 
tucopa, April, 3; Victorville, April, May, 3; 7 mi. east of Walker Pass, 
April, 5; Yermo, April, May, 5. Utah: State label only, 1. 

MEXICO: Baja California [Norte?]: Metero Beach, April, 3. 

Remarks. The color of the head in auriculata varies geographically. In 
Arizona, Baja California, and most of southern California the vertex is 
orange, usually with a black stripe on the midline. In western Fresno 



SYSTEM ATICS AND PHYLOGENY 169 

County, eastern Kern County, and eastern Los Angeles County there is 
an evident reduction in the average size of the orange area. West of 
these areas the trend continues into the Coastal Range, where the orange 
area is commonly reduced to an inconspicuous spot on each of the 
tempora or is obliterated entirely. Individuals without orange on the 
vertex occur in western Los Angeles County at Griffith Park, Mint 
Canyon, and Neenach. In Ventura County the vertex is entirely black in 
11 of 14 specimens examined. At Frazier Park and nearby Lebec, in 
extreme southern Kern County, it is black in 61 (79 per cent) of 77 
specimens. From Santa Barbara County five specimens are available 
(San Marcus Pass and Ventucopa), all with the orange area reduced 
to a small spot at the tempora. The single specimen from San Luis Obispo 
County (Morro Bay) is similar. Northward, in San Benito County, two 
of ten specimens from Pinnacles have the vertex entirely black. At San 
Lucas, Monterey County, it is entirely black in 3 of 4 specimens ex- 
amined; at Carmel, Monterey County, in 33 (75 per cent) of 44 speci- 
mens; and at Santa Cruz, Santa Cruz County, in 1 of 2 specimens. 

Correlated with the reduction of the orange area of the vertex in 
auriculata from the Coastal Range area of California there is a tendency 
for the color of the elytra to be darker and less metallic than is usual 
elsewhere in the range of the species. As far as I can tell, however, there 
is no comparable tendency for darkening of the pubescence. 

Included among the material studied in the collection of the San 
Jose State College is a male of auriculata found mated with a female of 
sublaevis at Coalinga, California. This apparently unique case of inter- 
specific mating of Meloidae was previously reported by Edwards ( 1951 ) . 

The first instar larva of auriculata has been described by MacSwain 
(1956). The type is in the collection of the Academy of Natural Sciences 
of Philadelphia. 

Bionomics. The following food plants have been recorded, primarily in 
California: African daisy, Encelia, and Geraea canescens (Compositae); 
Argemone and Eschscholtzia ( Papaveraceae ) ; Cuscuta utnbellata (Con- 
volvulaceae ) ; Dicentra ( Fumariaceae ) ; Eriodictyon and Phacelia (Hy- 
drophyllaceae ) ; Lupinus (Leguminosae); Pentstemon spectabilis (Scro- 
phulariaceae ) ; Salvia (Labiatae); Sambucus ( Caprifoliaceae ) ; Sphae- 
ralcea and S. ambigua (Malvaceae); and Verbena ( Verbenaceae ) . 

Lytta (Poreospasta) nigripilis (Fall) 

Cantharis nigripilis Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 303. 
Lytta nigripilis, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 91, 
pi. 14. 

Differs from auriculata as follows. 

Head usually black with a small orange frontal spot; in one female 



170 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

from Potwisha, Tulare County, California, top of vertex dark orange. 
Elytra dark metallic blue or blue-black, only rarely with a tinge of green. 
Pubescence uniformly piceous, or at most with the tips of the pubescence 
of the under surface of the thorax colorless. 

Male. Antennae (Fig. 100) with distal segments as a rule a little more 
moniliform and more noticeably wide than preceding segments. Pale 
pubescence ( pad ) of first segment of hind tarsi frequently limited to less 
than apical half of segment, never to more than half. Genitalia as in 
Figure 333; gonoforceps less elongate; gonostyli more robust, the mesal 
hooks located at or basad of midpoint between apex and base of gono- 
styli; aedeagus with dorsal margin more or less evenly rounded apically 
and with dorsal hook more massive, not strongly recurved. 

Type locality. Sierra Nevada of California, from Tulare County to 
Tuolumne County. 

Geographic distribution. Sierra Nevada and northern coastal region of 
California. (See Fig. 44.) 

Seasonal distribution. March — July 10. 

Records. UNITED STATES: California: State label only, 2; Ash 
Mountain, April, June, 3; Bass Lake, June, 10; Big Trees, Sequoia Na- 
tional Park, June, 1; 3 mi. south of Camino, June, 2; Camp Nelson, Tulare 
County, July, 1; Clear Lake, May, 2; Clearlake Oaks, May, 9; Colfax, 
May, 11; Colony Mill Road, Tulare County, June, 4; Comfort, Mendocino 
County, 1; Coulterville, April, 7; Glenn ville, April, 1; Havilah, June, 1; 
Hobergs, May, 2; Kern County, March, 3; Kaweah, 4; Malpitas, 3; Mari- 
posa County, May, June, 3; Meadow Valley, Plumas County, June, 1; 
Mendocino County, June, 11; Miami Ranger Station, Madera County, 
June, 5; Oakhurst, June, 1; Placerville, April, 2; Potwisha, Tulare County, 
June, 40; Riverton, July, 4; Seneca, June, 2; Sequoia National Park, 2000 
ft.-3000 ft., April, May, 60; Shut Eye Mountain, Madera County, June, 7; 
Snowline Camp, Eldorado County, June, July, 49; Springville, June, 5; 
Strawberry, June, 2; Sugar Pine, 4; Tuolumne County, May, 6; Wawona, 
June, 3. 

Remarks. The species appears to have a disjunct distribution, with 
a small population in the northern Coastal Range of California isolated 
by the Central Valley from the main population in the Sierra Nevada. 
There seems to be no morphological difference between the two popula- 
tions. 

The range of nigripilis is strictly allopatric with respect to that of 
auriculata. Along the coast of California there is apparently a sizable 
gap between the ranges of the two forms. In southern California the main 
nigripilis population extends to the southern end of the Sierra Nevada, 
which cuts deeply into the range of auriculata. Apparently in this region 



SYSTEMATICS AND PHYLOGENY 171 

nigripilis cannot exist in the dry lowlands, while auriculata is not suc- 
cessful in the higher, more mesic habitat of nigripilis. 

Since nigripilis and auriculata are very closely related, their allopatric 
distributional relationship suggests that they may ultimately prove to 
be only racially distinct. However, I have not found any definite indica- 
tion that they are capable of interbreeding. On the basis of the characters 
of the male genitalia they are 100 per cent separable. Some overlap be- 
tween them occurs in all other characters, but the variation producing this 
overlap tends in the main to be sporadic and nongeographic in nature, 
with no discernible tendency for the two species to converge morpho- 
logically where they approach each other geographically. Variation in 
head color in auriculata constitutes an exception in that it is clearly geo- 
graphical. However, here again there is no indication of intergradation 
between auriculata and nigripilis in a geographically intermediate area. 
It would therefore seem that the relatively poor differentiation of the 
two forms in most characters is a reflection of their close phylogenetic 
relationship rather than the result of introgression at the present time. 
Field investigations centered in the area of close geographic approxima- 
tion of the two species in Kern County will undoubtedly contribute 
greatly to a further clarification of the situation. 

The first instar larva of nigripilis has been described by MacSwain 
(1956) from specimens obtained from adults collected in Eldorado 
County, California. As to be expected, the larva differs from that of 
auriculata in only minor points. 

The type of nigripilis is in the Museum of Comparative Zoology. I 
have examined two cotypes, labeled southern California, in the United 
States National Museum. 

Bionomics. The following food plants have been recorded: Ceanothus 
(Rhamnaceae), Eriogonum ( Polygonaceae ) , Lupinus and Vicia (Legu- 
minosae), and Ranunculaceae. 

Lytta (Poreospasta) hoppingi Wellman 

Lytta hoppingi Wellman, 1912, Ent. News, vol. 23, p. 35. Van Dyke, 1929, 
Bull. Brooklyn Ent. Soc., vol. 24, p. 131. 

Black. Head with an elongate orange or piceous frontal spot between 
eyes; tempora frequently orange or piceous also. Pronotum orange with a 
wide median stripe and anterior margin black; stripe scalloped at sides, 
sometimes narrowed toward base. Wings light brown, the anal region 
paler. Length: 11 mm.-13 mm. 

Head quadrate; vertex smooth, shiny, rather finely, moderately densely 
punctate; frontal area a little more densely punctate, irregularly, shallowly 
impressed above and below frontal spot, rarely slightly alutaceous; pubes- 



172 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

cence rather long, erect. Eyes moderately wide, scarcely emarginate. 
Pronotum elongate, slender, one-tenth to three-tenths longer than wide; 
sides rather evenly arcuate from base to apex, without indication of lateral 
angles; disk weakly convex; surface and pubescence as on vertex. Scutel- 
lum subtruncate. Elytra noticeably broadened apically; surface weakly 
alutaceous, subimpunctate; pubescence in basal region long, as on head 
and pronotum, becoming shorter but not sparser toward apex. Outer hind 
tibial spur not or only slightly wider than inner spur, very obliquely 
truncate; inner spur obtuse. Tarsal pads weakly developed, similar to 
those of lugens. Tarsal claws weakly curved. 

Male. Antennae ( Fig. 101 ) reaching base of pronotum or surpassing 
it by one segment; segments elongate moniliform, III-X subequal in 
width or with distal segments very slightly wider; X noticeably shorter 
than rest. Fifth abdominal sternum entire. Emargination of sixth sternum 
(Fig. 256) triangular. Genitalia as in Figure 334. 

Female. Antennae (Fig. 142) reaching one segment beyond middle of 
pronotum; segments III-VI subequal in width; VII-X becoming progres- 
sively wider. Sixth abdominal sternum truncate. 

Type locality. Coalinga, California. 

Geographic distribution. Foothills at the southern end of the Central 
Valley of California. ( See Fig. 46. ) 

Seasonal distribution. March 21 — June. 

Records. UNITED STATES: California: Coalinga, May, 10 (including 
holotype and one paratype); Delano, March, 2; Tulare County, April, 2; 
Visalia, March, June, 17. 

Remarks. Morphologically hoppingi is quite similar to lugens. Its color 
pattern is unique for the genus Lytta. 

The type of hoppingi is in the United States National Museum (ex- 
amined ) . 

Lytta (Poreospasta) lugens (LeConte) 

Cantharis lugens LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 161. Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. 112; 1883, Trans. 
Amer. Ent. Soc., vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. Soc., vol. 27, 
p. 299. 

Lytta lugens, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 355. 

Black, with or without a tiny piceous or orange frontal spot between 
eyes. Wings light brown with colorless anal region. Length: 11 mm. -17 
mm. 

Head subquadrate; vertex smooth, shiny, moderately coarsely, moder- 
ately densely punctate, with a few very fine punctures interspersed with 



SYSTEMATICS AND PHYLOGENY 



173 



larger ones; frontal area more densely punctate, alutaceous, irregularly, 
shallowly impressed; pubescence moderately long, erect. Eyes moderately 
wide, scarcely emarginate. Pronotum (Fig. 186) elongate, slender, one- 
fifth longer than wide; sides straight and rather weakly divergent from 
base to before middle, then convergent; disk weakly convex; surface 
shiny, more coarsely and for the most part more densely punctate than 
vertex, generally distinctly alutaceous; pubescence as on vertex. Scutellum 
rounded. Elytra rugose, subimpunctate; pubescence very short, erect, 
sparse, with some longer setae at base and along margins. Outer hind 
tibial spur (Fig. 196) not or only very slightly wider than inner spur, 
distinctly shorter, flattened or very obliquely truncate; inner spur obtuse. 
Tarsal pads weakly developed, divided on fore and middle tarsi; pale 
pubescence ( pad ) completely absent on first segment of middle tarsi and 
all segments of hind tarsi. Tarsal claws moderately curved. 

Male. Antennae (Fig. 102) reaching base of pronotum; segments elon- 
gate moniliform; III— VI subequal in width; VII-X slightly wider; III and 
X shorter than rest ( except II ) ; XI about three-fourths as long as IX and 
X combined. Fifth abdominal sternum very shallowly, broadly emargi- 
nate. Emargination of sixth sternum (Fig. 257) triangular. Genitalia as 
in Figure 335; aedeagus with a single ventral hook. 




Fie. 45 (left). Distribution of Lytta sublaevis. 

Fig. 46 (center). Distribution of Lytta lugens (circles) and Lytta hoppingi (squares). 



Fig. 47 (right). Distribute 
( squares ) . 



.ytta aeneipcnnis (circles) and Lytta refulgent- 



174 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Female. Antennae (Fig. 143) reaching middle of pronotum; segments 
III-V subequal in width; VII-X becoming progressively wider. Sixth 
abdominal sternum truncate or weakly notched. 

Type locality. San Diego, California. 

Geographic distribution. Southwestern California, from Los Angeles 
and San Bernardino counties to San Diego County. (See Fig. 46.) 

Seasonal distribution. May 27 — July. 

Records. UNITED STATES: California: State label only, 2; Barton 
Flats, San Bernardino County, July, 2; Corona, 1; Coronado, May, 1; Los 
Angeles, 1; Pinon Flat, San Jacinto Mountains, May, June, 7; Ribbon- 
wood, 1; San Diego, 1; Vandevanter Flat, San Jacinto Mountains, June, 1. 

Remarks. This species may be immediately separated from other all- 
black species of Lytta by its elongate pronotum and slender outer hind 
tibial spurs. It is most similar structurally to hoppingi. 

The type of lugens is in the Museum of Comparative Zoology. 

Bionomics. Specimens have been collected in the San Jacinto Moun- 
tains on Argemone ( Papaveraceae ) , Eriogonum ( Polygonaceae ) , and 
Lupinus ( Leguminosae ) . 

Lytta (Poreospasta) aeneipennis (LeConte) 

Cantharis aeneipennis LeConte, 1851, Ann. Lye. Nat. Hist. New York, 
vol. 5, p. 160. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 113; 1883, 
Trans. Amer. Ent. Soc., vol. 10, p. 311. Fall, 1901, Occas. Papers California 
Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc., vol. 27, p. 299. 

Lytta aeneipennis, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 
6, p. 334. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 91. 

Vertex and median part of frontal area orange; rest of head black. 
Pronotum orange, finely margined with black; a short, brownish streak 
often present on midline before middle. Elytra metallic blue-green, green, 
or blue. Under surface and femora dark metallic blue or blue-black. 
Scutellum, tibiae, and tarsi black. Wings pale brown, the base nearly 
colorless. Pubescence piceous, with a sericeous luster; setae long and 
erect except on elytra, giving beetle a hairy appearance. Length: 5 mm.- 
12 mm. 

Head quadrate or nearly so; tempora not prominent; dorsal margin 
weakly rounded; vertex moderately shiny, finely granular, finely, sparsely 
to moderately densely punctate; where densest, punctures separated by 
an average distance greater than the diameter of a single puncture; frontal 
area more densely punctate, finely, weakly alutaceous, with a smooth, 
impunctate area at center; pubescence fine, about two-thirds as long as 
fourth segment of middle tarsus. Eyes small. Pronotum transverse, one- 
tenth to (rarely) one-fourth wider than long; outline varying between 



SYSTEMATICS AND PHYLOGENY 175 

the extremes shown in Figures 187 and 188; disk almost always with a 
shallow impression on each side behind middle; surface a little alutaceous, 
otherwise as on vertex and with similar pubescence; basal impressed line 
weak; basal margin not strongly inflexed. Scutellum truncate. Elytra ru- 
gose, impunctate; some long, erect setae at base, but surface otherwise 
with only scattered, short, subrecumbent setae. Pubescence on under sur- 
face, coxae, trochanters, and femora fine, long, as on head. Outer hind 
tibial spur moderately slender, not more than twice as wide as inner 
spur, acute at apex; inner spur acute (Fig. 198). Tarsal pads weakly 
developed, undivided; first segment of middle and hind tarsi entirely 
piceous pubescent. Tarsal claws slender. 

Male. Antennae (Fig. 103) reaching or slightly surpassing base of 
pronotum; distal segments about twice as long as wide; IV-X subequal 
in length and width; XI shorter than IX and X combined. Fifth abdominal 
sternum very shallowly emarginate. Sixth sternum variable in shape 
(Figs. 258-59), shallowly emarginate; a fine median groove running from 
posterior margin to base of sternum. Genitalia as in Figure 336; aedeagus 
with a single ventral hook. 

Female. Antennae (Fig. 144) not reaching base of pronotum. Fifth 
abdominal sternum entire. Sixth sternum truncate or feebly emarginate, 
lacking a distinct median groove. 

Type locality. Santa Ysabel, California. 

Geographic distribution. Southwestern California, from San Luis Obispo 
County to San Diego County. ( See Fig. 47. ) 

Seasonal distribution. March 18 — May. 

Records. UNITED STATES: California: State label only, 21; near 
Cariso [Carrizo] Creek (Horn, 1873) (not mapped); 4 mi. west of Coa- 
huila [Coachella], April, 89; 10 mi. southeast of Creston, March, 1; El- 
sinore, 3; The Gavilan, Riverside County, April, 1; Pomona, 4; Poway, 1; 
Riverside, April, 4; San Diego, 1; Santa Isabel [Ysabel] (LeConte, 1851); 
Simmler, March, April, 29; Ventucopa, April, 6. 

Remarks. This species most closely resembles refulgens structurally. 
It is easily distinguished from all related species in having both the 
vertex and the pronotum orange in color. 

The larva has been described by MacSwain (1956). The type is in the 
Museum of Comparative Zoology. 

Bionomics. The series from near Coahuila was collected on Eschscholt- 
zia ( Papaveraceae ) , as previously reported by MacSwain (1956). 

Lytta (Poreospasta) refulgens Horn 

Lytta refulgens Horn, 1870, Trans. Amer. Ent. Soc, vol. 3, p. 91. MacSwain, 
1956, Univ. California Publ. Ent., vol. 12, p. 93. 



176 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Ltjtta 

Cantharis refulgens, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 114; 
1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. 
Soc., vol. 27, p. 299. 

Brilliant metallic green or bluish green. Head with a moderately large 
orange frontal spot. Pronotum orange, finely margined with green or 
black and with a large metallic green or bluish green median spot which 
is connected either to anterior margin or (rarely) basal margin. Elytra 
with a brassy luster. Tibiae brassy or black. Tarsi black. Wings darker 
than in either aeneipennis or crotchi. Pubescence either piceous or pale; 
setae long and erect except on elytra, giving beetle a hairy appearance. 
Length: 7.0 mm.-11.5 mm. 

Head quadrate or nearly so; tempora not prominent; dorsal margin 
weakly rounded; vertex smooth, shiny, moderately coarsely, sparsely 
punctate; punctures separated by an average distance twice or nearly 
twice as great as the diameter of a single puncture; frontal area weakly 
alutaceous, more densely punctate, with an impunctate area at center; 
pubescence fine, varying in different specimens from less than half as 
long to fully as long as fourth segment of middle tarsus; as a rule the 
pubescence is distinctly shorter than in aeneipennis and crotchi. Eyes 
small, straight on margin facing antennal insertion. Pronotum transverse, 
one-tenth to one-fourth wider than long, more or less suboval in outline; 
sides not strongly curved; disk usually with a shallow impression on each 
side behind middle; surface weakly alutaceous, not granular, rather 
finely, weakly, very sparsely punctate, the center of disk nearly impunc- 
tate; pubescence as on vertex; green discal spot one-third width of pro- 
notum; basal impressed line deep; basal margin strongly reflexed. Scutel- 
lum rounded. Elytra rugose, impunctate; some long, erect setae at base, 
but surface otherwise with only scattered, short, subrecumbent setae. 
Pubescence on under surface, coxae, trochanters, and femora fine, sparse, 
as long as on head, not as hairy as in aeneipennis and crotchi. Middle 
tibiae weakly bowed. Outer hind tibial spur slender, slightly wider 
than to (rarely) nearly twice as wide as inner spur, very obliquely ex- 
cavate, acute at apex; inner spur acute. Tarsal pads weakly developed, 
undivided on at least fore and middle tarsi. Tarsal claws slender. 

Male. Antennae as in Figure 104, reaching or extending one or two 
segments beyond base of pronotum. Fifth abdominal sternum entire. 
Emargination of sixth sternum shallow, as in typical chloris and some 
aeneipennis; sternum flattened at center but without a median groove. 
Genitalia as in Figure 337; aedeagus with a single ventral hook. 

Female. Antennae ( Fig. 145 ) a little longer than in aeneipennis, reach- 
ing base of pronotum. Middle tibiae more strongly bowed than in male. 
Sixth abdominal sternum truncate, not flattened. 



SYSTEMATICS AND PHYLOGENY 177 

Type locality. Millerton, California. Millerton was formerly a com- 
munity in northern Fresno County, near Friant. Its site is now covered by 
Millerton Lake. 

Geographic distribution. Western Sierran foothills from Fresno County 
to northern Kern County, California. (See Fig. 47.) 

Seasonal distribution. March 30 — April 24. 

Records. UNITED STATES: California: State label only, 2; Glennville, 
March, 1; 6 mi. west of Kernville, April, 4; Millerton (Horn, 1870); 
Walker Pass, Kern County, April, 12. 

Remarks. The first instar larva of this species has been described by 
MacSwain (1956). The type is in the collection of the Academy of Natu- 
ral Sciences of Philadelphia. 

Bionomics. Adults have been recorded from Ceanothus (Rhamnaceae) 
and Lupinus ( Leguminosae ) (MacSwain, 1956). 

Lytta (Poreospasta) crotchi (Horn) 

Cantharis Crotchii Horn, 1874, Trans. Amer. Ent. Soc, vol. 5, p. 38; 1883, 
Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Trans. Amer. Ent. Soc, 
vol. 27, p. 299. 

Lytta crotchi, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 95. 

Dark metallic green or greenish blue, often slightly brassy. Head with 
a small, round orange frontal spot. Pronotum orange, finely margined 
with black. Antennae, tibiae, and tarsi black. Pubescence piceous, with- 
out conspicuous silver reflection, in greater part very long, erect, giving 
beetle a very hairy appearance. Length: 6 mm. -10 mm. 

Head shape as in aeneipennis; vertex more distinctly microreticulate 
than in that species, rough, coarsely, deeply, densely punctate; punctures 
separated by an average distance less than the diameter of a single punc- 
ture; frontal area more densely punctate, some of the punctures con- 
fluent; entire head conspicuously clothed with fine pubescence which 
is about as long as fourth segment of middle tarsus. Pronotum transverse, 
one-tenth to one-half longer than wide, oval, with sides more strongly 
convergent at apex than at base; disk finely granular, like vertex, but less 
coarsely, less densely, and much more shallowly punctate, most of the 
punctures confined to sides, the center of disk very sparsely punctate; 
pubescence like that of head; basal impressed line deep; basal margin 
strongly reflexed. Elytra rugose, impunctate, sparsely, regularly, conspicu- 
ously clothed with long, fine, erect setae which are fully as long as or even 
longer than those of head. Wings similar to those of aeneipennis but 
darker brown. Pubescence on under surface of coxae, trochanters, and 
femora long, fine, like that of elytra. Abdomen less densely punctate and 
pubescent than in aeneipennis. Tarsal pads weakly developed, distinctly 



178 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

divided on all segments, the midline glabrous; pale pubescence of first 
segment of hind tarsi limited to apex. Tarsal claws weakly curved. 

Male. Antennae (Fig. 105) surpassing base of pronotum by two or 
three segments; segments elongate, not so closely articulated or so filiform 
as in aeneipennis; distal segments at most twice as long as wide. Fifth and 
sixth abdominal sterna as in aeneipennis except that sixth sternum some- 
times lacks an impression and fine groove on midline. Genitalia as in 
Figure 338; aedeagus with two ventral hooks. 

Female. Antennae reaching base of pronotum. Middle tibiae more 
strongly curved than in male. Sixth abdominal sternum truncate, not 
impressed or grooved. 

Type locality. San Diego, California. 

Geographic distribution. Southwestern California, from Los Angeles 
County to San Diego County. (See Fig. 48.) 

Seasonal distribution. March 8 — April 18. 

Records. UNITED STATES: California: State label only, 2; 4 mi. 
west of Coahuila, April, 3; Pine Valley, San Diego County, April, 95; 
San Diego (Horn, 1874); San Dimas, Dalton Canyon, Los Angeles 
County, April, 3; Tanbark Flat, Los Angeles County, April, 8. 

Remarks. This is an unusually hairy species resembling refulgens in 
color. It differs most conspicuously from refulgens and the related but 
differently colored aeneipennis by the presence of long pubescence over 
the entire surface of the elytra; in this respect it is like comans. It also 
differs from these two species in having two (rather than one) ventral 
hooks on the aedeagus. Structurally, crotchi is most similar to aeneipennis. 

The first instar larva of crotchi has been described by MacSwain 
( 1956 ) . The type is in the collection of the Academy of Natural Sciences 
of Philadelphia. 

Bionomics. Adults have been collected on Ceanothus (Rhamnaceae) 
at Tanbark Flat and on Eschscholtzia ( Papaveraceae ) near Coahuila. 
An interesting ecological relationship between crotchi and aeneipennis 
is suggested by the fact that these species occur sympatrically, are active 
as adults at the same time, and share at least one food plant ( Eschscholt- 
zia) in common. 

Lytta (Poreospasta) rathvoni LeConte 

Lytta rathvoni LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 335; 1857, in Reports of explorations and surveys . . . Mississippi River to 
the Pacific Ocean, vol. 12, book 2, pt. 3, p. 21. MacSwain, 1956, Univ. Cali- 
fornia Publ. Ent, vol. 12, p. 96. 

Cantharis rathvoni, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 112; 
1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 1901, Occas. Papers Cali- 
fornia Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc, vol. 27, p. 299. 



SYSTEMATICS AND PHYLOGENY 179 

Brilliant metallic green, brassy green, or brass-colored, with the usual 
orange frontal spot. Elytra metallic violet or greenish violet, varying 
occasionally to a nearly pure green. Pubescence colorless. Length: 8 mm.- 
17 mm.; average with standard error (50 specimens), 12.1 mm. ± .3 mm. 

Differs structurally from the Sierran foothills race of chloris as follows. 

Pubescence longer, denser, and comparatively conspicuous on under 
surface. Microreticulation of elytra less distinct. Male antennal segments 
slightly more elongate ( Fig. 107 ) ; segment V 2.4-3.2 ( average, 25 speci- 
mens, 2.73) times as long as wide. Male sixth abdominal sternum (Fig. 
261) rather deeply impressed medianly and more deeply triangularly 
emarginate. Female sixth sternum (Fig. 283) emarginate medianly. 

Type locality. Near Sacramento, California. 

Geographic distribution. California, from the Sierran foothills in 
Tulare County north to Calaveras and Sacramento counties. (See Fig. 
48.) 

Seasonal distribution. March 17 — June 14. 

Records. UNITED STATES: California: El Mirador, April, 1; El 
Portal, May, 1; Fresno, May, June, 10; Kaweali, 3; Lemoncove, March, 
66; Millerton (Horn, 1873); Mokelumne Hill, 1; Potwisha, March, May, 4; 
near Sacramento (LeConte, 1853); Salt Creek, Tulare County, April, 1; 
Sequoia National Park, April, May, 43; Springville, June, 2; Squaw Valley, 
Fresno County, April, 6. 

Remarks. The only species with which rathvoni is likely to be confused 
is chloris, which occurs with it in Tulare County; see remarks for chloris. 

The first instar larva of rathvoni has been described by MacSwain 
(1956). The type is in the Museum of Comparative Zoology. 

Bionomics. Most of the specimens I have seen from Lemoncove were 
collected by J. W. MacSwain, who has recorded them ( MacSwain, 1956 ) 
from Lupinus ( Leguminosae ) and Eschscholtzia ( Papaveraceae ) . 

Lytta (Poreospasta) chloris (Fall) 

Cantharis chloris Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 303. 
Lytta chloris, Linsley and MacSwain, 1942, Amer. Midland Nat., vol. 27, 
p. 406. 

Lytta chloris ?, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 92. 

Brilliant brassy green; the development of the brassy quality varies 
considerably but as a rule is strongest on the head and pronotum. Head 
with a moderate-sized, oval or diamond-shaped orange frontal spot. 
Pubescence entirely colorless (Sierran foothills) or entirely piceous, or 
(rarely) with individual setae piceous basally and colorless apically; 
length of pubescence variable. Length: 7 mm.-14 mm.; average with 
standard error (75 specimens), 10.1 mm. ± .2 mm. 



180 BIONOMICS, SYSTEM ATICS, AND PHYLOGENY OF Lyttd 

Head quadrate to triangular; surface micropunctate, usually distinctly 
microreticulate and satiny in texture, not glistening, moderately coarsely, 
rather sparsely punctate; pubescence sparse, short to moderately long. 
Pronotum subhexagonal, as wide as to nearly one-fifth wider than long; 
disk uneven; midline finely impressed, at least at center; surface usually 
even more distinctly micropunctate than head, usually transversely alu- 
taceous, moderately coarsely, rather sparsely, irregularly punctate; pu- 
bescence as on vertex. Elytra rugose, very distinctly microreticulate, 
impunctate; pubescence consisting of some very short setae inconspicu- 
ously scattered over surface except at immediate base (rarely as much 
as basal third ) and margins, where the setae are as long as those on pro- 
notum. Pubescence on under surface sparse, variable in length. Legs 
slender, unmodified. Outer hind tibial spur twice as wide as inner spur, 
acute; inner spur acute. Tarsal pads moderately well developed, un- 
divided on fore and middle tarsi; ventral pale pubescence of first segment 
of hind tarsi limited to apex when clothing setae of tarsi are piceous. 
Tarsal claws weakly curved. 

Male. Antennae (Fig. 106) very slender, reaching three or four seg- 
ments beyond base of pronotum; segments more nearly cylindrical than 
in stijgica and comans; segment V 2.1-3.1 (average, 28 specimens, 2.49) 
times as long as wide. Fifth abdominal sternum very shallowly emargi- 
nate. Sixth sternum typically only slightly impressed and very shallowly, 
obtusely emarginate, as in Figure 262. Genitalia as in Figure 339; aedea- 
gus with only a single ventral hook (Sierran foothills) or with a small 
second hook also developed. 

Female. Antennae (Fig. 146) not reaching base of pronotum. Sixth 
abdominal sternum either entire or weakly emarginate medianly. 

Type locality. Kern County or Tulare County, California. 

Geographic distribution. California. There appear to be two separate 
populations. One of these ranges through the valleys and foothills of the 
Coastal Range from Mount Diablo, Alameda County, south to the Cu- 
yama Valley and western Kern County (vicinity of Wasco). The other 
occupies the lower slopes and foothills of the Sierra Nevada in Tulare and 
Kern counties. (See Fig. 49.) 

Seasonal distribution. March 14 — May 16. 

Records. UNITED STATES: California: Adobe Creek, Stanislaus 
County, April, 4; Arvin, March, 12; 20 mi. east of Bakersfield, March, 1; 
2 mi. northeast of Caliente Mountain, San Luis Obispo County, March, 
2; Coalinga, March, 4; Cottonwood Creek, Kern County, (ambiguous; 
not mapped) March, 5; Cuyama Valley, Kern and Santa Barbara coun- 
ties, 6; Glenn ville, April, 55; Havilah, May, 1; Hospital Canyon, San 
Joaquin County, March, 22; Kaweah, 6; Kern County, March, 1; La 



SYSTEMATICS AND PHYLOGENY 



181 



Panza, April, 3; Lemoncove, March, 57; Monterey County, March, 1; 
Mount Diablo, April, 1; Mount Hamilton, April, 3; North Fork Kaweah 
River, April, 1; San Antonio Valley, 35 mi. south of Livermore, April, 54; 
San Jose, April, 9; Santa Clara County, April, 1; Sequoia National Park, 
3000 ft.-5000 ft., April, May, 9; Silver Creek, Santa Clara County, April, 8; 
Simmler, March, 21; Tulare County, 4; Visalia, March, 1; 14 mi. west 
of Wasco, April, 3; Woody, April, 8. 

Remarks. Throughout the range of chloris, the head, pronotum, and 
elytra are usually very distinctly microreticulate and the male sixth ab- 
dominal sternum is usually relatively flat and very shallowly, evenly 
emarginate. These two characters are of great assistance in separating 
chloris from stygica and comans, and the second one is very useful in 
separating chloris and rathvoni. However, there is sufficient variation 
in chloris, as well as in the other species, that these characters must be 
used in conjunction with others. 

Typical chloris, as described by Fall, seems to be confined to the 
Sierran foothills in Tulare and Kern counties. If we exclude for the 
moment two short series of specimens from Arvin and Cottonwood Creek, 
this population may be characterized as having the pubescence colorless, 
the aedeagus with a single ventral hook, and the female sixth abdominal 
sternum almost always entire (exceptions in two specimens from Glenn- 
ville and one from Lemoncove). In the Coastal Range area to the west, 




Fig. 48 (left). Distribution of Lytta crotchi (circles) and Lijtta rathvoni ( squares ] 
Fig. 49 (center). Distribution of Lytta chloris. 
Fig. 50 (right). Distribution of Lytta comans. 



182 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

and including the Cuyama Valley and western Kern County, chloris is 
represented by a population which differs from the typical population in 
having the pubescence piceous, the aedeagus almost always with two 
ventral hooks ( exceptions in one specimen each from Coalinga and Simm- 
ler), and the female sixth sternum entire in only 55 per cent of the 
specimens. 

In view of the generally low level of morphological differentiation 
among species in the Stygica Group, the Sierran foothills and Coastal 
Range populations of chloris might reasonably be treated as separate 
species on the basis of the characters given above. However, in the 
absence of information as to whether they can exist in sympatry, I prefer 
at present to regard them as races of a single species; this is the inter- 
pretation suggested by MacSwain (1956) in first calling attention to the 
Coastal Range population in connection with his description of its first 
instar larva. It is to be hoped that with further field work the apparent 
distributional gap between the two populations will be filled, thus per- 
mitting an objective determination of whether intrinsic isolating mecha- 
nisms between them exist. 

The species with which chloris is likely to be confused are rathvoni, 
stygica, and comans. The Sierran foothills race of chloris occurs sym- 
patrically with rathvoni in Tulare County and with stygica in Kern 
County. In the southern part of its range the Coastal Range race of 
chloris is sympatric with southern stygica in the Cuyama Valley; to the 
north it occurs just inland of the coastal population of stygica. It occurs 
in broad sympatry with comans. In distinguishing between chloris and 
these species, it is convenient to treat the two races separately. 

From rathvoni, the Sierran foothills race of chloris differs by the fol- 
lowing combination of characters: average size smaller; elytra brassy green 
(not purple); microreticulation of elytra slightly more distinct; pubes- 
cence on under surface shorter and sparser, that on abdomen generally 
very short and inconspicuous; male antennal segments a little less elongate; 
male sixth abdominal sternum flatter ( not as deeply impressed ) and more 
shallowly, evenly emarginate; female sixth sternum usually entire. Since 
chloris and rathvoni maintain their separate identities in their area of 
sympatry, they fall within the usual definition of species. There is, how- 
ever, a tendency for them to vary toward each other in a large series 
of both species from Lemoncove. In this series the vast majority of 
specimens are typical of either chloris or of rathvoni, but a few are diffi- 
cult to assign to either one or the other species. The nature and extent 
of the differences separating typical chloris and rathvoni are such that 
the intergradation at Lemoncove is quite likely the result of individual 
intraspecific variation, but the possibility of some introgression between 
them should not be completely discounted. 



SYSTEM ATICS AND PHYLOGENY 183 

From sympatric stygica, the Sierran foothills race of chloris differs 
as follows: color brassy green (green, blue, or black in stygica); pubes- 
cence colorless; head and pronotum usually distinctly microreticulate 
( smooth or nearly so in most stygica ) ; microreticulation of elytra slightly 
more distinct; male sixth abdominal sternum usually flatter and more 
shallowly and evenly emarginate; female sixth sternum usually entire. 
In a male from Arvin the pubescence is entirely piceous, while in 13 other 
specimens from this locality and in a series of 4 specimens from Cotton- 
wood Creek (Kern County; exact location not known) at least part of 
the pubescence is piceous with only the tip colorless. The entirely 
piceous-pubescent Arvin male is also atypical of chloris in having a 
small second hook indicated on the ventral side of the aedeagus. In addi- 
tion, three (of a total of ten) of the females represented in the two 
series have the sixth abdominal sternum definitely emarginate medianly. 
On the basis of these characters, these series might be regarded as inter- 
grades with the Coastal Range race of chloris, but an equally likely 
hypothesis is that their atypical features are the result of introgression 
from stygica, which has been taken at these same localities. In support 
of this second hypothesis there is the fact that the microreticulation of 
the head and pronotum is weak in both series. It must be admitted, how- 
ever, that this evidence is tenuous in view of the wide range of normal 
variation in this character in chloris. Here again, as in the case of sym- 
patric chloris and rathvoni in Tulare County, field studies of the popula- 
tions involved seem to be essential for a real understanding of what 
appears to be a very complex variational problem. 

The Coastal Range race of chloris differs from stygica by the same 
characters as does the Sierran foothills race except that the color of the 
pubescence is the same (entirely piceous) in both and the value of 
the form of the female sixth abdominal sternum as a basis for separation 
is much lessened inasmuch as it is emarginate (as in stygica) in about 
half the females of chloris in the Coastal Range area. Additional char- 
acters for separation are the presence of a second ventral hook on the 
aedeagus of Coastal Range chloris and a rather distinct difference in male 
antennal length (segments more elongate in chloris) between chloris 
and the northern coastal population of stygica. 

From comans, Coastal Range chloris differs in the same characters that 
distinguish it from northern coastal stygica except that there is no per- 
ceptible difference in the color of the integument ( green in both species ) . 
There is one additional and very useful difference, however, and that is 
the presence in comans of long, erect setae on the entire sutural half of 
the elytral surface. Chloris usually has some moderately long or long 
setae at the immediate base of the elytra and along the margin. In occa- 
sional specimens these are present as far back as the basal third of the 



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SYSTEM ATICS AND PHYLOGENY 185 

elytra, but I have seen no specimens of chloris that could be confused 
with comans on the basis of this character. 

For a more concise comparison of the characters of chloris, stygica, 
comans, and rathvoni than that provided by the above discussion, the 
reader is referred to Table V. 

The type of chloris is in the Museum of Comparative Zoology. 

Bionomics. Linsley and MacSwain (1942) found chloris present in 7 
(.92 per cent) of 759 cells examined at a nesting site of Anthophora 
linsleyi Timberlake 20 mi. east of Bakersfield, Kern County. Food plants 
recorded for the adult beetles (one or two records each) are Baerio, B. 
chrysostoma, and Loyia ( Compositae ) . 

Lytta (Poreosposta) stygica (LeConte) 

Cantharis stygica LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 161. Horn, 1873, Proe. Amer. Philos. Soc., vol. 13, p. 113; 1883, Trans. 
Amer. Ent. Soc., vol. 10, p. 311. Fall, 1901, Oecas. Papers California Acad. 
Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc., vol. 27, pp. 293-94, 299. 

Cantharis smaragdula LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 

5, p. 160. New synonymy. 

Lytta stygica, LeConte, 1853, Proe. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 335; 1857, in Reports of explorations and surveys . . . Mississippi River to 
the Pacific Ocean, vol. 12, book 2, pt. 3, p. 21. 

Lytta smaragdula, LeConte, 1853, Proe. Acad. Nat. Sci. Philadelphia, vol. 

6, p. 334; 1857, in Reports of explorations and surveys . . . Mississippi River 
to the Pacific Ocean, vol. 12, book 2, pt. 3, p. 21. MacSwain, 1956, Univ. 
California Publ. Ent., vol. 12, p. 93. 

Lytta dolosa LeConte, 1861, Proe. Acad. Nat. Sci. Philadelphia, p. 354. 

Cantharis stolida Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 302. New 
synonymy. 

Cantharis purpurescens Fall, 1901, Trans. Amer. Ent. Soc., vol. 27, p. 302. 
New synonymy. 

Cantharis smaragdina [sic], Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 
294 (lapsus calami) . 

Cantharis difficilis Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 303. Neiv 
synonymy. 

Lytta arborea Wellman, 1912, Ent. News, vol. 23, p. 34. New synonymy. 

Lytta purpurescens, Linsley and MacSwain, 1942, Amer. Midland Nat., 
vol. 27, p. 406, fig. 7. 

Lytta nevadensis Van Dyke, 1947, Pan-Pacific Ent., vol. 23, p. 155. New 
synonymy. 

Lytta stolida, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 94. 

Varying from bright metallic green or blue to a pure black. Head with 
a small orange frontal spot. Pubescence piceous. Length: 7 mm. -15 mm. 

Head usually quadrate, rarely triangular; surface usually smooth, rarely 
mircroreticulate; surface sparsely, finely punctate; pubescence sparse, 
moderately long. Pronotum varying from quadrate to subhexagonal; sur- 
face and pubescence as on head. Elytra rugose, distinctly microreticulate, 



186 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

impunctate; pubescence short. Outer hind tibial spur twice as wide as 
inner spur, acute; inner spur acute. Tarsal pads moderately well de- 
veloped, undivided on fore and middle tarsi; ventral pale pubescence 
of first segment of hind tarsi limited to apex. Tarsal claws as in Fig- 
ure 202. 

Male. Antennae varying in length between extremes shown in Figures 
108 and 109; see also Table VI. Fifth abdominal sternum very shallowly 
emarginate or entire. Emargination of sixth sternum shallowly triangular 
( Fig. 263 ) . Genitalia as in Figure 340; aedeagus almost always with only 
a single ventral hook, rarely with a weak second hook. 

Female. Antennae as in Figure 147 (northern California) or with seg- 
ments more elongate than shown. Sixth abdominal sternum shallowly 
emarginate. 

Type locality. Of stygica, Oregon; of smaragdula, mountains around 
Santa Ysabel, California; of dolosa, Mendocino, California; of stolida, 
San Francisco, California; of purpurescens, Pasadena or Riverside, Cali- 
fornia; of difficilis, San Diego, California; of arborea, near Hamburg, 
Siskiyou County, California (see remarks, below). 

Geographic distribution. (See Fig. 51.) Beginning in southern Idaho 
the range of this species extends northwest (probably discontinuously ) 
to the Cascade Mountains in Washington and thence south along these 
mountains and the Coastal Range to northern California, where it is 
divided into two arms by the Central Valley of California. One arm con- 
tinues south along the humid coast, passes through the San Francisco 
Bay region, and terminates in the Santa Cruz Mountains. The other arm 
extends south through the Sierra Nevada to Tuolumne County. South of 
the Santa Cruz Mountains and south of Tuolumne County there is hiatus 
of range of considerably more than 100 miles. In eastern California the 
range commences again in the Inyo, Panamint, and Argus mountains, 
Inyo County; crosses the southern end of the Sierra Nevada and the 
Tehachapi mountains; and continues south, mainly along the San Gabriel, 
San Bernardino, San Jacinto, and Santa Rosa mountains and associated 
foothills, to the Coastal Range of San Diego County. To the east popula- 
tions are known from the Spring Mountains in southwestern Nevada and 
from an area in southeastern Arizona extending from the Tucson region 
to the White Mountains. 

Notwithstanding the fact that it occurs at some low, arid localities 
in the southern half of its range, stygica is clearly unable to make a 
deep penetration into true desert habitat as shown by its avoidance of 
the Great Basin, the more arid portions of the Central Valley of Cali- 
fornia, and all but the western margins of the Mojave and Colorado 
deserts. The populations in the Argus Mountains and the Spring Moun- 



SYSTEMATICS AND PHYLOGENY 



187 



tains seem to be isolated from each other and from other populations. 
The same may be true of the populations in the Panamint Mountains 
and Inyo Mountains, and its probably true for the population in south- 
eastern Arizona. Presumably these populations are relicts surviving from 
a time (most likely the last Pluvial Period) when less severe climatic 
conditions permitted the range of the species to extend more or less 
continuously east from the southern Sierras of California into Arizona. 
If this is true, then we can expect to find additional isolated populations 
in southern Nevada and northern Arizona. 

The distributional gap between the northern and southern segments 
of the range of stygica is discussed below. 

Seasonal distribution. March 8 — July 21. There are relatively few 
March records in the northern half of the range and relatively few June 
records in the southern half. July records are rare throughout the range. 




Fig. 51. Distribution of Lytta stygica. Letters are used to identify samples in Table VI. 



188 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Records. 3 UNITED STATES: Arizona: State label only, 2; Cortaro, 14; 
Globe, 1; San Carlos, Gila River, 2; Tucson, 32; White Mountains, 1. 
California: Alameda County: Arroyo Mocho, 5 mi. south of Livermore, 
14; Berkeley, 1; Leona Hills, 1; hills back of Oakland, 25; Oakland, 2. Al- 
pine County: Hope Valley, 20; Lake Alpine, 4. Amador County: Ten Rise 
[?] (not located), 1. Butte County: Jarboe Pass, 11. Eldorado County: 
County label only, 1; Echo Lake, 2; Glen Alpine Creek, 1; Meyers, 6300 
ft., 12; Placerville, 1; Pollock Pines, 2. Humboldt County: County label 
only (Nunenmacher material, probably from Weitchpec), 9; Garberville, 
1; Fort Seward, 1. Inyo County: Argus Mountains, 38; Lone Pine, 1; 
Mazurka Canyon, Inyo Mountains, 2; Mountain Spring Canyon, Argus 
Mountains, 6; Panamint Mountains, 4; Westgard Pass Plateau, 3. Kern 
County: 20 mi. east of Bakersfield, 9; 18 mi. south of Bakersfield, 7; Fort 
Tejon, 1; Glennville, 3; Kernville and 5 mi. north of Kernville, 24; Havilah, 
9; Lebec, 4000 ft., 4; Piute Mountains, 5; Rand, 3; Searles Station, 4; 
"South. Cala.," 5 (including coptype of purpurescens); Tehachapi, 45; 
Walker Pass and 3 mi. east of Walker Pass, 97; Willow Springs, 2. Lake 
County: Hullville, 1; Kelseyville, 1; Middletown, 5. Los Angeles County: 
County label only, 6 (including cotype of purpurescens); Chicago, 1; 
Fairmont, 3; Gorman, 1; Lancaster, 5; Little Rock, 1; Los Angeles, 2; 
Mint Canyon, 1; Palmdale and 2 mi. east of Palmdale, 11; Pasadena (Fall, 
1901, p. 185); Pearblossom and 6 mi. east of Pearblossom, 13. Marin 
County: County label only, 4; Fairfax, 12; Kentfield, 1; Mill Valley, 1; 
Muir Woods, 5; Ross, 1; San Francisco (including Mission Hills, Point 
Lobos, and Twin Peaks), 21; Sausalito, 1. Mendocino County: County 
label only, 1; Baechtel Creek, 3 mi. west of Willits, 1; Eagle's Nest, (not 
located) 1; Mendocino (LeConte, 1861); Pygmy Forest, Fort Bragg, 16; 
Rancheria Creek, near Boonville, 3; Ryan Creek, 57; 2 mi. north of Wil- 
lits, 1. Modoc County: Badger Well, 3; Davis Creek, 4800 ft., 1; Hacka- 
more, 1. Napa County: Calistoga, 25; Mount Saint Helena, 54; Pope 
Valley, 5. Nevada County: County label only, 1; Chicago Park, 2; Nevada 
City, 7. Placer County: County label only, 2; Colfax, 10; Dutch Flat, 1; 
Emigrant Gap, 6. Plumas County: County label only, 2; Johnsville, 2; 
Meadow Valley, 4000 ft.-5000 ft., 2; Onion Valley, 2; 4 mi. west of Quincy, 
2. Riverside County: 4 mi. west of Coahuila [= Coachella], 20; Idyllwild, 
4; Indio, 2; Marion Mountain Camp, San Jacinto Mountains, 1; Palm 
Springs, 1; Riverside (Fall, 1901, p. 185); Santa Rosa Mountains, 5; 10 
mi. southeast of Temecula, 1. San Bernardino County: County label only, 
7; Barstow, 1; Barton Flats, 1; Deep Creek, 17; Forest Home, 4; Joshua 
Tree National Monument (Bell Picnic Area), 2; Twenty-nine Palms, 3; 

3 In order to save space, months in which specimens have been collected are 
omitted. In the case of California I have departed from the system used elsewhere 
in this paper in listing the localities by counties. 



SYSTEM ATICS AND PHYLOGENY 189 

Victorville, 3. San Diego County: County label only, 1; Borego, 3; Borego 
State Park, 1; Dulzura, 2800 ft., 1; Mason Valley, 1; San Diego, 1; moun- 
tains near Santa Ysabel (LeConte, 1853). San Mateo: Brisbane, 6; Half 
Moon Bay, 2; Kings Mountain Road, 3; Peseadero, 6; Portola State Park, 
1; San Andreas Lake, 2; San Mateo, 2. Santa Barbara: Cuyama River, 6. 
Santa Cruz County: Ben Lomand, 4; Big Basin, 6; Boulder Creek, 6; 
between Boulder Creek and Big Basin, 1; Santa Cruz, 2. Shasta County: 
Black Mountain, 1; 5 mi. east of Burney, 1; Castella, 34; Lassen Na- 
tional Park (Kelley's Resort and Manzanita Lake), 4. Sierra County: 
Gold Lake, 7. Siskiyou County: County label only (Nunenmacher mater- 
ial, including type of arborea, presumably from near Hamburg), 5; 
Dunsmuir, 31; Hamburg, 1; McCloud, 7; Scott Bar, 4; Yreka, 1. Solano 
County: 5 mi. south of Dixon, 4. Sonoma County: Trinity, 8. Trinity 
County: County label only, 1; Big Flat, 1; Carrville, 2400 ft.-2500 ft., 11; 
Coffee Creek, 1; Eagle Creek, 1; Hayfork, 3000 ft., 3; Plummer Springs 
Ranger Station (not located), 1; Trinity River Camp (not located), 1. 
Tulare County: Lemoncove, 5; Sequoia National Park, 5000 ft.-7000 ft., 2. 
Tuolumne County: Long Barn, 2; Strawberry, 1. Idaho: Boise, 2692 
ft., 4 ; Caldwell, 2375 ft., 2; Emmett, 2373 ft., 2; Payette, 7; Pine, 12; 
Pocatello, 3. Nevada: Kyle Canyon, Charleston Mountains [Spring 
Mountains], 17 (including type of nevadensis). Oregon: Alsea, 3; Ash- 
land, 7; 20 mi. east of Cascadia, 1; Corvallis, 16; Giles Lake, Portland, 1; 
Gold Hill, 1; Homestead, 1; Jacksonville, 1; Kane Creek (not located), 2; 
Keno, 4; Klamath County, 5; 10 mi. northwest of Klamath Falls, 1; 
Langden Lake, 2; Mount Jefferson, 2; North Santiam River, 10 mi. north 
of highway 20, Linn County, 2; Prospect, 7; Warner Mountains, 4; Yon- 
calla, 1. Washington: Dryden, 1; Easton, 1: Ellensburg, 1; Lake Wanna, 
1; Rebels Ridge, Pomeroy, 1; Walla Walla, 10. 

Remarks. As presently defined, stygica includes a highly variable com- 
plex of populations. In the past authors have attempted to distinguish 
a number of species within the complex, but I do not believe that this 
procedure is justified on the basis of the data now available. 

Stygica varies in color from bright metallic green or blue through 
shades of deep blue to a pure, nonmetallic black. Pure black specimens 
seem to be relatively scarce, although in many of the dark specimens 
the metallic luster is visible only in the proper light or under magnifica- 
tion. The metallic coloration seems to be entirely structural in origin. 
When "black" specimens are immersed in a liquid, their luster is con- 
siderably enhanced. 

North of California and east of the Cascade Mountains all specimens 
examined are either bright green or blue-green. In the Cascades of Wash- 
ington they are darkened to a deep blue or blue-black. To the south, in 
an area encompassing the Coastal Range and Cascades of Oregon and 



190 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

extending south into California through northern Siskiyou County and 
along the Coastal Range, the metallic coloration is, as a general rule, 
either poorly developed or absent. This does not apply, however, to 
specimens from the less humid, inland side of the Coastal Range of Cali- 
fornia. Thus in Lake and Napa counties there is a pronounced tendency 
for the development of green coloration, and even the darkest specimens 
have some metallic luster. Specimens from near Dixon Springs, in Sonoma 
County, well within the Central Valley, are bright green. In San Mateo 
County, south of San Francisco, specimens from Brisbane, Half Moon 
Bay, and Pescadero are black, while those from localities farther inland 
( San Andreas Lake, Portola State Park, and Kings Mountain ) are bright 
green. Finally, in Santa Cruz County 5 specimens examined are entirely 
black, 12 are greenish black, and 3 are green. 

Returning to northern California, the material from Modoc County is 
either black or dark blue. In the northern Sierras, the Mount Shasta 
region, and Trinity County the color is consistently a bright metallic 
green or blue-green. This same bright coloration is characteristic also 
of all specimens of stygica from Arizona and southern Nevada and all 
those from southeastern California as far west as Tulare County, extreme 
eastern Kern County (Walker Pass), the San Bernardino and San Jacinto 
mountains, and the Coastal Range in San Diego County. In the southern 
Sierran foothills of central Kern County, on the Tehachapi Mountains, 
and in extreme northern Los Angeles County (Fairmont and Lancaster) 
the great majority of specimens available are black with a brassy or 
purplish luster developed to a very high degree, while the green speci- 
mens tend to be quite dark. In the Tehachapi series, which is the only 
large one from the general area, there is complete intergradation from 
essentially pure black to bright metallic green. To the south, all speci- 
mens from Pearblossom and Palmdale, Los Angeles County, are bright 
green, as are six specimens from the Cuyama Valley, Santa Barbara 
County. Further south, 2 of 4 specimens from Forest Home, San Ber- 
nardino County, are black; the single specimen from near Temecula, 
western Riverside County, is metallic black; and, finally, the two speci- 
mens from southwestern San Diego County (San Diego and Dulzura) 
are pure black. 

There is marked geographic variation in antennal length in stygica. 
In general the antennae are relatively elongate in the southern part of 
the range of the species and short in the northern part. This variation 
is expressed graphically in Table VI, using the proportion of length to 
width of the male fifth segment as an index of relative antennal length. 

Along the northern part of the coast of California and particularly 
in the San Francisco Bay region, there is a tendency for the head to be 
more triangular in shape than usual. The pronotum varies considerably 



SYSTEM ATICS AND PHYLOGENY 191 

in proportion of length to width and in shape but without clear-cut geo- 
graphic pattern. The aedeagus is also notably variable. As a general rule, 
only a single hook is present, but occasional specimens from Alpine and 
Kern counties, California, have a weakly developed second hook. 

Morphological differences between first instar larvae of stygica from 
eastern Kern County ( = smaragdida ) and the San Francisco Bay region 
(= stolida) have been described by MacSwain (1956). 

For a summary of the principal differences between adults of stygica 
and the closely related species chloris, rathvoni, and comans, see Table V. 

The types of stygica, smaragdida, dolosa, stolida, purpurescens, and 
difficilis are in the Museum of Comparative Zoology. The type of neva- 
densis is in the collection of the California Academy of Sciences (Type 
no. 5866, examined); the type series is labeled as collected in June, not 
July as stated by Van Dyke ( 1947 ) . 

Wellman's (1912) description of arborea was based on an unspecified 
number of specimens collected at Weitchpec, Humboldt County, Cali- 
fornia, and near Hamburg, Siskiyou County, California, May 20, 1911 
and June 2, 1911, respectively, by F. W. Nunenmacher. The type locality 
was given as Humboldt County, and it was stated that the type was in 
the Wellman collection. There is, however, no specimen labeled as the 
type in the Wellman collection in the United States National Museum. 
On the contrary, one of several Siskiyou County specimens in the Nunen- 

Table VI. Variation in the Proportions ( Length/ Width ) of the 
Male Fifth Antennal Segment in Lytta stygica* 



Washington, norlheostern Oregon, and Idaho (A) , 

Western Oregon ond northern coostol and 

extreme northern California (B) l_ 

Central coostol California (C) 

North-central California and the 

Sierra Nevada (D) 

Inyo Mountoins and Panammt Mountains, i 

California (E) , I i (N=6) 




Mountains, Nevada (F) 



California (G) 



Northwestern Son Bernardino County ond 
norlheostern Kern County, California (H) 

Southern Kern County, Los Angeles County, ond 
west-central San Bernardino County, Cal (I) 

Southern Son Bemordino County, Riverside County, 
ond Son Diego County, California (J) 



Barbara County, Colifornio (K) 
rn Arizona (U 




a Observed limits of variation shown by length of horizontal lines; means marked by narrow 
vertical triangles; bars represent one standard deviation on each side of mean; black portions 
of bars represent two standard errors of the mean on each side of the mean. 

Letters are used to identify samples on the map shown in Figure 51. 



192 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

macher collection in the Chicago Natural History Museum bears a hand- 
printed label that is unquestionably Wellman's, designating it as the 
"type" of "Lytta aborea [sic] Wellm." Another of the Siskiyou County 
specimens in the same collection has Wellman's cotype label (with the 
specific name again misspelled). Whatever the cause of this gross incon- 
sistency on Wellman's part, his labeling of the Siskiyou County specimen 
as the type of arborea must surely take precedence over his published 
statement. 

Bionomics. Linsley and MacSwain (1942) found stygica (as pur- 
purescens) infesting 29 (3.82 per cent) of 759 cells of Anthophora lins- 
leyi Timberlake examined from a nesting site 20 mi. east of Bakersfield, 
Kern County, California. This same nesting site was also parasitized by 
the closely related chloris and by tenebrosa. 

In the adult stage stygica feeds on the pollen of a variety of spring 
flowers, including some domesticated ones. At present there is no indica- 
tion of food plant differences between the various geographic segments 
of the species population. The food plants recorded are as follows: African 
daisy [ArctotisP], Layia, and Rafinesquia neomexicana ( Compositae ) ; 
Lesquerella gordani ( Crucif erae ) ; Arctostaphylos and Rhododendron 
(Ericaceae); Eriodictyon calif ornicum and Phacelia ( Hydrophyllaceae ) ; 
Iris ( Irideaceae ) ; Salvia (Linsley and MacSwain, 1942) (Labiatae); As- 
tragalus and Lupinus (several records) ( Leguminosae ) ; Eschscholtzia 
and E. calif ornica (several records) ( Papaveraceae ) ; Ceanothus (Rham- 
naceae); Cercocarpus, Cowania stansburiana, and Primus (Rosaceae); 
Orthocarpus ( Scrophulariaceae ) ; and Lycium (Solanaceae). 

Lytta (Poreospasta) comans, new species 

Brilliant green or brassy green. Pubescence piceous. Length: 7 mm.- 
13 mm.; average with standard error (79 specimens), 10.1 mm. ± .2 mm. 

Structurally like the northern Coastal Range population of stygica 
except as follows: Clothed throughout with long, erect pubescence. Male 
antennal segments somewhat more elongate and cylindrical in shape; 
segment V 1.8-2.5 (average, 50 specimens, 2.26) times as long as wide. 

Geographic distribution. Coastal Range and western side of the Central 
Valley, California, from Yolo County south to the Cuyama Valley. (See 
Fig. 50.) 

Seasonal distribution. February 27 — June 25. 

Type material. Holotype male and allotype female from Los Altos, 
Santa Clara County, California, March 1937, E. S. Ross, in the collection 
of the California Academy of Sciences. 

Additional records. UNITED STATES: California: Adobe Creek, Stan- 
islaus County, March, 46; Alma, 2; Alum Rock Park, Santa Clara County, 



SYSTEMATICS AND PHYLOGENY 193 

April, 1; Arroyo Bayo, Mount Hamilton, April, 4; Big Sur, Monterey 
County, April, 1; Cache Creek Canyon, Yolo County, April, 1; Cedar 
Ridge, Alameda County, May, 1; Clayton, April, 1; Clear Creek, Cu- 
yama Canyon, March, 1; Contra Costa County, May, 2; Del Puerto Can- 
yon, March, 1; Guadalupe Lake, Santa Clara County, April, 2; Los Altos, 
March, April, 55; Monterey, March, 4; Mount Hamilton, May, 2; Niles 
Canyon, Alameda County, March, 2; Paraiso Springs, April, May, 10; 
Pine Canyon, Monterey County, March, 5; Pine Mountains, San Luis 
Obispo County, April, 1; Pinnacles National Monument, April, 9; Putah 
Canyon, Yolo County, March, April, 62; San Antonio Valley, Santa Clara 
County, April, 6; San Jose, April, 1; Somersville, March, 9; Stevens Creek, 
Santa Clara County, April, 1. 

Remarks. This species is a derivative of stygica, from which it is only 
weakly differentiated morphologically. In the northern half of its range 
the available data seem to indicate that it is an inland replacement of 
the northern Coastal Range population of stygica. In Alameda County, 
in the Livermore region, the two populations have been recorded only a 
few miles from each other. To the south, in the absence of stygica, comans 
is able to extend its range all the way to the coast. The southern limit 
of known range is at Clear Creek, a tributary of the Cuyama River. 
Very probably it enters the Cuyama Valley itself, and in that event it is 
probably in contact with the southern population of stygica, which enters 
the valley from the east. It is unfortunate, but the relationship between 
comans and stygica in this southern region cannot be determined ac- 
curately at the present time due to the lack of critical material. 

Throughout its range comans maintains its distinctiveness, and despite 
its obviously close phylogenetic relationship with stygica, it seems to 
represent a separate species. 

See remarks for chloris. 

Bionomics. There are one or two records each of adults collected on 
Ceanothus cuneatus (Rhamnaceae), Cryptantha (Boraginaceae), Erio- 
dictyon (Hydrophyllaceae), and Eschscholtzia calif ornica (Papave- 
raceae ) . 

MOERENS GROUP 

Head with a pale (orange) frontal spot. Head sparsely micropunctate, 
minutely granular, finely punctate. Eyes narrow. Male antennae reaching 
one to three segments beyond base of pronotum; intermediate segments 
often incrassate, so that they are as wide as or wider than distal segments. 
Male hind trochanters spined or angulate on ventral margin in most 
species (always so if intermediate antennal segments not incrassate). 
Middle tibiae strongly bowed. One or both spurs of male middle tibiae 



194 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

usually thickened and sticklike. Outer hind tibial spur thickened, ob- 
liquely truncate; inner spur slender, spiniform. Male fore and middle tarsi 
at least slightly heavier than in female. Tarsal pads well developed, dense, 
undivided; female with pale pubescence (pad) of first segment of fore 
and middle tarsi limited to apical half or less. Male fifth abdominal ster- 
num deeply emarginate. Male sixth sternum elongate; emargination very 
deeply, narrowly triangular; a lightly sclerotized area medianly (rarely 
absent in moerens), this area and lateral lobes of sternum clothed with 
long pubescence. Male genitalia with each gonostylus bearing a mesal 
hook and clothed ventrally with short but conspicuous setae, or if mesal 
hooks absent, then gonostyli clothed ventrally with long setae; aedeagus 
with a single ventral hook or (childi) none. 

Remarks. This group contains nine species, divided into three sub- 
groups. It is widely distributed in western North America north of 
Mexico, with two of the species (moerens and childi) ranging south 
onto the peninsula of Baja California and another (nuttalli) as far east 
as Minnesota (Fig. 8). 

Food plants have been recorded for six of the nine species of the 
group. These records suggest that the preferred, if not the only, food 
plants are species of Leguminosae. So far as known the beetles feed on 
both flowers and leaves. 

Key to Subgroups 

1. Black or very dark blue; first segment of male fore tarsi straight, 
not excavate at base; male fore tibiae sometimes with only a single 

spur each 2 

Bright metallic green, blue, purple, or violaceous; first segment of 
male fore tarsi bent, with a deep, socket-like excavation at base; male 
fore tibiae always with two well-developed spurs each; gonostyli of 
male genitalia lacking mesal hooks . . . Cyanipennis Subgroup ( p. 204 ) 

2. Gonostyli of male genitalia each with a mesal hook at apex; male 
antennal segment I not densely punctulate on ventral side; pronotum 
never marked with orange; male fore tibiae sometimes with posterior 

spur obsolete or very small Moerens Subgroup (p. 194) 

Gonostyli of male genitalia lacking mesal hooks; male antennal seg- 
ment I densely punctulate on ventral side; pronotum frequently in 
part orange; male fore tibiae with posterior spur always well devel- 
oped Childi Subgroup ( p. 215 ) 

Moerens Subgroup 

Entirely black or very dark blue; body sometimes with a metallic blue 
luster. Frontal spot one-fifth to one-fourth as wide as frontal area between 



SYSTEMATICS AND PHYLOGENY 195 

eyes. Male antennae with intermediate segments incrassate ( insperata 
and nigrocyanea ) or not; all segments but II longer than wide. Pronotum 
hexagonal or nearly so, wider than long; lateral lobes obtusely rounded; 
disk flattened, granular, rather dull. Male hind trochanters each angulate 
on ventral margin or armed with a sharp spine. Male middle tibiae each 
with one or both spurs somewhat modified. First segment of male fore 
tarsi cultriform but not distorted. Emargination of male fifth abdominal 
sternum triangular. Male genitalia with each gonostylus bearing a mesal 
hook at apex; gonostyli very sparsely clothed ventrally with fine, short 
setae; aedeagus with a single ventral hook (rudiment of a second hook 
sometimes present in moerens). 

Remarks. This subgroup includes the species moerens, navajo, in- 
sperata, and nigrocyanea. The first of these appears to be fairly common, 
ranging from Oregon south through California. The species insperata 
has been found only in southern California. Navajo is known from a 
single collection in northern Arizona, while nigrocyanea has been col- 
lected in western Colorado, eastern Utah, and Wyoming. 

Key to Species 

1. Male antennae clavate, with intermediate segments much narrower 
than distal segments; male hind trochanters angulate on ventral 

margin, not spined 2 

Male antennae with intermediate segments incrassate, at least in part 
as wide as or wider than distal segments; male hind trochanters each 
armed with a distinct spine on ventral margin 3 

2. Head and pronotum (especially latter) sparsely punctate; elytra 
finely, weakly rugose, moderately shiny; male abdominal sterna rela- 
tively smooth, shiny; lateral lobes of male sixth abdominal sternum 
not divergent (Fig. 264); male genitalia as in Figure 341; male pos- 
terior fore tibial spur often much reduced in size or absent 

moerens ( p. 196 ) 

Head and pronotum densely punctate; elytra coarsely, strongly ru- 
gose ( subreticulate ) , dull; male abdominal sterna finely rugose, 
dull; lateral lobes of male sixth abdominal sternum strongly divergent 
( Fig. 265 ) ; male genitalia as in Figure 342; male posterior fore tibial 
spur well developed navajo (p. 200) 

3. Entirely black; head sparsely short-pubescent; pronotum subglabrous; 
male fore tibiae each with two spurs; male middle tibiae not modified 
as below; male sixth abdominal sternum (Fig. 266) extremely elon- 
gate, with lateral lobes not divergent insperata (p. 201 ) 

At least elytra very dark blue; head and pronotum clothed with long, 
erect pubescence, hairy; male fore tibiae each with a single spur; 



196 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

male middle tibiae acutely produced at apex and with posterior spur 
projecting at a wide angle; male sixth abdominal sternum (Fig. 267) 
shorter, with lateral lobes strongly divergent .... nigrocyanea ( p. 202 ) 

Lytta (Poreospasta) moerens (LeConte) 

Cantharis moerens LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 5, 
p. 216. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 110; 1874, Trans. 
Amer. Ent. Soc, vol. 5, p. 39; Trans. Amer. Ent. Soc, vol. 10, p. 311. Fall, 
1901, Occas. Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. 
Ent. Soc, vol. 27, p. 298. 

Lytta moerens, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 332; 1857, in Reports of explorations and surveys . . . from the Mississippi 
River to the Pacific Ocean, vol. 12, book 2, pt. 3, p. 21. MacSwain, 1956, 
Univ. California Publ. Ent., vol. 12, p. 88, pi. 14. 

Cantharis incommoda Horn, 1883, Trans. Amer. Ent. Soc, vol. 10, p. 312. 
Fall, 1901, Trans. Amer. Ent. Soc, vol. 27, p. 298. New synonymy. 

Lytta nunenmacheri Wellman, 1912, Ent. News, vol. 23, p. 36. 

Lytta incommoda, Linsley and MacSwain, 1942, Univ. California Publ. Ent., 
vol. 7, p. 195, tbl. 

Black, with or without a metallic bluish luster. Wings variable in 
color, generally nearly colorless with light brown apex, sometimes color- 
less only in anal region. Pubescence piceous on upper surface of body, 
either piceous or nearly white on under surface. Length: 11 mm.-21 mm. 

Head as wide as or slightly wider than long, subtriangular to quadrate; 
surface moderately shiny, never particularly rough, sparsely to moder- 
ately densely punctate; pubescence long on sides and frontal area, shorter 
on vertex. Pronotum subhexagonal; disk irregularly impressed; surface 
more granular than on vertex, sparsely punctate before middle, somewhat 
more densely so and often finely alutaceous behind; pubescence as on 
vertex. Scutellum subtruncate or rounded. Elytra finely, weakly rugose, 
moderately shiny, impunctate, essentially glabrous. Outer hind tibial spur 
twice as wide as inner spur; truncature oval, almost always laminate. 
Tarsal claws ( Fig. 203 ) heavy, strongly curved. 

Male. Antennae (Fig. 110) clavate; segments IV-X subequal in length 
to III, becoming progressively much wider and more moniliform; X 
globular. Hind trochanters weakly angulate to almost spined along ven- 
tral margin distad of middle, the margin rather sharp. Anterior fore tibial 
spur sinuate, longer and heavier than posterior spur, which is often rudi- 
mentary or absent on one or both fore tibiae. Middle tibial spurs heavy, 
either both spiniform or posterior one flattened, sticklike. Abdominal 
sterna clothed with short setae, these much shorter and less conspicuous 
than in female; surface finely punctate, finely granular but shiny. Emargi- 
nation of fifth abdominal sternum broad, ridged or notched at apex. 
Emargination of sixth sternum as in Figure 264; sternum usually with a 



SYSTEMATICS AND PHYLOGENY 



197 



lightly sclerotized median area; lateral lobes of sternum not divergent, 
less densely pubescent than in related species. Genitalia as in Figure 
341; gonoforceps robust; gonostyli not bowed in ventral view; setae 
longer and less sparse than in navajo; aedeagus as shown or with a rudi- 
ment of a second hook basad of apical one. 

Female. Antennae (Fig. 148) reaching a little beyond middle of pro- 
notum, more strongly clavate than in insperata, with segment IV three- 
fifths as wide as IX; intermediate segments more moniliform than in male. 
Sixth abdominal sternum (Fig. 284) broadly, shallowly emarginate. 

Type locality. Of moerens, Sacramento, California; of incommoda, 
southern California; of nunenmacheri, Orleans Bar, Humboldt County, 
California. 

Geographic distribution. Widespread in California, extending north 
into western Oregon. ( See Fig. 52. ) Occurs in both mountains and low- 
lands. 

Seasonal distribution. March 4 — July 10. 

Records. UNITED STATES: California: State label only, 21; Antioch, 
April, May, 7; Arvin, March, April, 33; Bairs Ranch, Redwood Creek, 
Humboldt County, June, 1; 18 mi. north of Bakersfield, April, 1; Berkeley, 
April, 4; Bass Lake, 2500 ft., June, 2; 10 mi. north of Blackwells Corner, 




Fig. 52 ( left ) . Distribution of Lytta moerens. 

Fig. 53 (right). Distribution of Lytta navajo (circle), Lytta insperata (squares), and 
Lytta nigrocyanea ( triangles ) . 



198 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttO 

April, 82; 5 mi. east of Carmel, May, 1; Carrville, 2400 ft.-2500 ft., May, 
June, 9; Coffee Creek, Trinity County, May, 4; Coloma, May, 2; Coro- 
nado, 8; Cypress Point, June, 1; Davis Creek, 4800 ft., Modoc County, 
June, 3; Delano, March, 9; El Portal, May, 1; Fresno, June, 2; Glenn ville, 
May, 1; Hullville, June, 4; Imperial County, April, 5; Indian Flat, [Guard 
Station], Mariposa County, June, 1; Indian Wells, Owens Valley (not 
located), May, 1; Kern County, March, 6; Kettleman City, May, 4; 
Lancaster, May, 4; Lanes Bridge, Fresno County, April, 43; Lone Pine, 
Inyo County, April-June, 29; Los Angeles County, April, July, 10; Lucerne 
Valley, 30 mi. south of Barstow, May, 62; McGee Creek, Mono County, 
June, 5; 20 mi. northwest of McKittrick, April, 2; near Mather, July, 70; 
Meadow Valley, 3500 ft.-4000 ft., Plumas County, June, 1; Milham City 
(not located), April, 112; Mojave, June, 2; Montara (Linsley and Mac- 
Swain, 1942); Muroc Dry Lake, San Bernardino County, April, 3; New- 
port Bay, May, 1; [Orleans Bar], Humboldt County, May, 12 (lectotype 
and paratypes of ntmenmacheri) ; Owens Valley, 4 (3 in May); Plumas 
County, June, 12; Point Reyes, May, 18; Putah Canyon [Creek], April, 1; 
Quincy and 4 mi. west of Quincy, May-June, 75; Rancho de Oso, Santa 
Cruz County, July, 3; Sacramento ( LeConte, 1851 ) ; Salida, March, 10; 
San Bernardino, May, 7; San Diego, 5; San Diego County, 3; San Fran- 
cisco, March, 3; San Jose, March, 2; San Luis Obispo, 3; Shasta County, 
July, 1; Snow Line Camp, El Dorado County (not located), June, 1; Tuo- 
lumne County, 4 ( 2 in July ) ; near Whitmore Hot Springs, Mono Count)', 
June, 1; Willits and 2 mi. north of Willits, May, 15. Nevada: State label 
only, 1. Oregon: State label only, 5; Coburg, July, 49; Corvallis, May-July, 
6; Independence, July, 1; Klamath Falls, June, 10; Marys Peak, May, 1; 
Monmouth, June, 2; Monroe, June, 2; Salem, June, 3. 

Remarks. The population of moerens south of the Central Valley of 
California and along the Owens River Valley east of the Sierra Nevada 
is clearly differentiated from the rest of the species population in con- 
sistently lacking the posterior spur on both fore tibiae of the male. In 
one of the specimens from Los Angeles County and in another from 
San Diego a small socket is present at the point where the missing spur 
should arise. In these specimens the socket has a continuous membranous 
covering, indicating that the absence of the spur is not the result of 
breakage. In all other males from the southern area there is no indica- 
tion of such a spur socket. Elsewhere within the range of moerens the 
posterior spur is absent from both fore tibiae only in two of a long 
series of males from Lanes Bridge, Fresno County, California. In a few 
additional males from Fresno County and a few from Plumas County, 
California, it is lacking on one fore tibia only. In all other males it is 
present on both fore tibiae although varying greatly in size. On the 
average, the spur is about half the length of the anterior fore tibial spur. 



SYSTEM ATICS AND PHYLOGENY 199 

At its smallest it is reduced to a short peg which does not extend beyond 
the base of the tibia and is therefore difficult to see unless the tibia is 
properly oriented under the microscope and examined rather closely. 
Variation in the size of the spur does not seem to be correlated with 
geographic distribution within the northern segment of the moerens 
population. 

Color in moerens varies clinally in a north-south direction. In Oregon 
and northern California almost all specimens are a pure black in color, 
the few exceptional ones being faintly metallic. Progressing southward, 
through central California, the metallic coloration occurs more frequently 
and becomes more obvious, the elytra tending to develop a purplish tone 
and the rest of the body a blue or green luster. South of the Central 
Valley of California and along the Owens River Valley the metallic luster 
is very conspicuously developed in all specimens. 

For the most part the outer hind tibial spur of moerens is quite constant 
in size and shape. However, in nearly a third of the specimens examined 
from Arvin, Kern County, California, as well as in four specimens from 
Oregon, it is much more strongly expanded than usual, much less ob- 
liquely truncate, and not at all laminate. 

LeConte based his original description and subsequent redescription 
( 1853) of moerens on a unique type. This specimen is now in the LeConte 
collection at the Museum of Comparative Zoology (no. 5125), where it 
was examined at my request by F. G. Werner. According to Dr. Werner 
(in lift.), the specimen is a female lacking any metallic luster. It is labeled 
with a gold disc, which indicates California, and another label with the 
abbreviation "Sac." [Sacramento] and Rathvon's name. 

Horn (1873 and subsequently), who based his definition of moerens 
on specimens from the southern part of the range, wrongly assumed that 
the metallic coloration and absence of the male posterior fore tibial spur 
characteristic of moerens in the southern area were constant features of 
the entire species. Subsequently, he encountered specimens of more 
northern moerens which he failed to recognize as conspecific with his 
"moerens" and which he described under the name incommoda. The type 
of incommoda is in the Horn collection at the Academy of Natural Sci- 
ences of Philadelphia. 

Lytta nunenmacheri has long been recognized as a synonym of in- 
commoda. I have examined 15 specimens from the type series of nunen- 
macheri, including a male and female of moerens (mounted in copulo) 
marked "Type." These last specimens, in the Chicago Natural History 
Museum (Nunenmacher collection), are labeled Humboldt County, 
California, May 22, 1911, collected by F. W. Nunenmacher. As it is not 
evident which of the two Wellman considered to be the type of nunen- 
macheri, selection of a lectotype is required, and I hereby designate the 



200 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

male of the pair as such. It should be pointed out that three of the speci- 
mens of Wellman's type series are representatives of blaisdelli rather 
than moerens, which accounts for some of the discrepancy between his 
description of nunenmacheri and the actual characters of moerens. In 
particular, his description of the antennae of nunenmacheri seems to 
apply more closely to blaisdelli than to moerens. 

Bionomics. The known larval hosts of moerens are Anthophora stan- 
fordiana Cresson (Linsley and MacSwain, 1942) and Colletes fulgidus 
Swenk (MacSwain, 1956). The former record is from a nesting site at 
Montara, California; the latter is from an unspecified locality. 

Adults of moerens have been collected on the following Leguminosae: 
lima beans at Independence, Oregon; Lupinus at Arvin, 5 mi. east of 
Burney, Indian Well, Lone Pine, Milham City (MacSwain, 1956), Muroc 
Dry Lake, and 4 mi. west of Quincy, all in California; and Vicia at Willits 
and 4 mi. north of Willits, California (MacSwain, 1956). In addition, 
there is a questionable record of radishes (Cruciferae) at Coburg, Ore- 
gon. 

Lytta (Poreospasta) navajo Werner 
Lytta navajo Werner, 1951, Psyche, vol. 57, p. 134, figs. 2, 4. 

Black with a metallic bluish luster. Wings nearly colorless. Pubescence 
piceous. Length: 16 mm. -23 mm. (Werner, 1951). 

Head strongly triangular, one-fifth wider than long; tempora well 
marked; surface moderately shiny, rough, densely, deeply punctate; pu- 
bescence long on sides and frontal area, shorter on vertex. Pronotum 
subhexagonal, one-seventh wider than long; disk irregularly impressed, 
uneven; surface as on vertex but finely alutaceous except for a smooth 
median area, much more densely punctate than in moerens, the sides 
very densely punctate; pubescence as on vertex. Scutellum subtruncate. 
Elytra coarsely, strongly rugose ( subreticulate ) , dull, impunctate, essen- 
tially glabrous. Outer hind tibial spur twice as wide as inner spur; trun- 
cature broadly oval, laminate. Tarsal claws heavy, strongly curved. 

Male. Antennae as in moerens. Hind trochanters subangulate along 
ventral margin distad of middle, the margin rather sharp. Anterior fore 
tibial spur sinuate, a little longer than posterior spur, which is well 
developed. Middle tibial spurs heavy, flattened, sticklike. Abdominal 
sterna clothed with short setae; surface finely punctate, rugose, dull. 
Emargination of fifth abdominal sternum as in moerens. Emargination of 
sixth sternum as in Figure 265; lateral lobes of sternum divergent, more 
densely, coarsely pubescent than in moerens. Genitalia as in Figure 342; 
gonoforceps more elongate than in moerens; setae of gonostyli extremely 



SYSTEMATICS AND PHYLOGENY 201 

short; mesal hook of each gonostylus not turned mesad; ventral hook of 
aedeagus obsolescent. 

Female. Not seen. Probably with sexual characters very similar to 
those of moerens. 

Type locality. 22 mi. north of Cameron, Coconino County, Arizona. 

Geographic distribution. Known only from the type locality, in north 
central Arizona. (See Fig. 53.) 

Seasonal distribution. The type series was collected May 19. 

Records. UNITED STATES: Arizona: 22 mi. north of Cameron, May, 
1 (paratype). 

Remarks. Werner (1951) described this species from a series of two 
males and three females, all bearing the same data. The type, a male, 
is in the Museum of Comparative Zoology (no. 28501), as is the allotype. 
The specimen I have seen is a male paratype in the collection of F. G. 
Werner. 

Bionomics. The type series was found feeding on Astragalus (Legu- 
minosae). 

Lytta (Poreospasta) insperata (Horn) 

Cantharis insperatus Horn, 1874, Trans. Amer. Ent. Soc, vol. 5, p. 39. 

Cantharis insperata, Horn, 1883, Trans. Amer. Ent. Soc, vol. 10, p. 311. 
Fall, 1901, Occas. Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. 
Amer. Ent. Soc, vol. 27, p. 298. 

Black. Wings light brown, with anal region almost colorless. Pubes- 
cence silvery on under surface of thorax, piceous elsewhere. Length: 
18 mm. -23 mm. 

Head as wide as long; sides rounded, moderately divergent above eyes; 
tempora rather prominent; surface moderately shiny, smooth, sparsely 
punctate; pubescence short, the setae on lower frontal area longer than 
rest. Pronotum subhexagonal, barely wider than long; disk deeply, 
broadly impressed at middle, shallowly impressed at sides behind middle; 
midline distinctly impressed for basal two-thirds; surface not alutaceous, 
rather dull, more sparsely, irregularly punctate than head, subglabrous. 
Scutellum rounded, distinctly foveo-sulcate. Elytra finely, weakly rugose, 
impunctate, essentially glabrous. Outer hind tibial spur (Fig. 197) four 
times as wide as inner spur; truncature nearly circular, not at all laminate. 
Tarsal claws rather weakly curved. 

Male. Antennae (Fig. Ill) with segments elongate filiform, slightly 
compressed, becoming somewhat moniliform, more rounded on lateral 
side than on mesal, subequal in width; IV-IX equal in length, nearly one- 
fifth longer than III; X as long as III, XI not quite two-thirds as long 
as IX and X combined, cylindrical for basal two-thirds, then tapered. Hind 



202 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

trochanters (Fig. 162) truncate at apex, armed with a spine on ventral 
margin near apex; ventral margin otherwise nearly straight. Anterior 
fore tibial spur straight, longer than posterior spur, which is well devel- 
oped. Posterior margin of middle tibiae produced at apex into a pointed 
plate which is short of middle of posterior spur; posterior middle tibial 
spur sticklike, twice as wide as anterior spur, projecting at a wide angle, 
truncate at apex. Emargination of fifth abdominal sternum (Fig. 218) 
moderately narrow, not notched at apex, lined with long, silky pubes- 
cence. Sixth sternum (Fig. 266) extremely elongate, cleft medianly for 
apical two-thirds; lightly sclerotized median area narrow, extending from 
basal fifth to apical fifth of sternum. Genitalia as in Figure 343; gono- 
forceps elongate, slender, tapered; aedeagus slender, with ventral hook 
well developed. 

Female. Antennae (Fig. 149) less strongly clavate than in moerens, 
with segment IV four-fifths as wide as IX. Sixth abdominal sternum entire, 
not at all emarginate. 

Type locality. Mojave Desert, California. 

Geographic distribution. Southern California from Ventura County 
to San Diego. (See Fig. 53.) 

Seasonal distribution. Specimens examined were collected May 26. 
Recorded by Fall at San Diego in April. 

Records. UNITED STATES: California: San Diego (Fall, 1901); 
Mojave Desert (Horn, 1874); Ventura County, May, 3. 

Remarks. Two of the specimens I have studied are in the collection of 
the California Academy of Sciences; the other specimen is in the collec- 
tion of the University of California. The type is in the collection of the 
Academy of Natural Sciences of Philadelphia. 

Rionomics. Essig's (1926) record of insperata as a pest of beets is 
highly doubtful in view of the evident rarity of the species. Essig mis- 
takenly figured specimens of Linsleya sphaeticollis (Say) as Lytta in- 
sperata, but this can hardly be the species in question either. Possibly 
the record refers to moerens. 

Lytta ( Poreospasta ) nigrocyanea Van Dyke 
Lytta nigrocyanea Van Dyke, 1929, Bull. Brooklyn Ent. Soc, vol. 24, p. 129. 

Black, with or without a dark blue luster. Elytra very dark blue or 
indigo, somewhat metallic. Wings nearly colorless. Pubescence piceous. 
Length: 19 mm. -20 mm. 

Head wider than long, strongly triangular; tempora prominent; dorsal 
margin straight; vertex, especially at tempora, distinctly tumid; surface 
rather dull, very densely punctate, the punctures becoming coarser and 
less dense on frontal area; surface extremely hairy, being densely clothed 



SYSTEMATICS AND PHYLOGENY 203 

with long, erect, silky pubescence. Pronotum hexagonal, wider than long; 
disk transversely impressed before middle, with a broad, somewhat 
foveate impression on each side at middle; surface rather dull, very 
densely punctate behind middle, more sparsely so before; pubescence as 
on vertex. Elytra finely rugose or rugose punctate, dull; pubescence sparse 
and very short except for some long setae at base. Outer hind tibial spur 
less than twice as wide as inner spur; truncature oval. Tarsal claws 
heavy, strongly curved. 

Male. Antennae with segments more elongate than in cyanipennis; IV- 
VIII subequal in length, incrassate, bulged apically on lateral side, tri- 
angular in shape; VII and VIII more strongly bulged and slightly 
wider than rest; IX and X shorter than preceding segments, as wide as 
VI. Hind trochanters truncate at apex, armed with a spine on ventral 
margin at distal third. Anterior fore tibial spur long, heavy, excavate 
basally; posterior spur obsolete. Apex of fore and middle tibiae acutely 
produced on posterior side. Posterior middle tibial spur sticklike, heavier 
than anterior spur, somewhat contorted. Emargination of fifth abdominal 
sternum broad, notched at apex; lateral lobes of sternum broadly rounded. 
Emargination of sixth sternum as in Figure 267; lateral lobes of sternum 
strongly divergent, curved ventrad at apex; lightly sclerotized median 
area triangular. Genitalia as in Figure 344; gonoforceps robust; gonostyli 
bowed in ventral view; aedeagus with ventral hook well developed. 

Female. Antennae clavate. Hind trochanters distinctly angulate on 
ventral margin at distal third. Fore tibiae each with two spurs. Apex of 
fore and middle tibiae less acutely produced than in male. Fifth ab- 
dominal sternum shallowly emarginate. Sixth sternum with a narrow, 
U-shaped emargination medianly, much as in childi. 

Type locality. Palisade, Colorado. 

Geographic distribution. Western Colorado, eastern Utah, and Wyom- 
ing. (See Fig. 53.) 

Seasonal distribution. The type series was collected May 7; the speci- 
mens from Utah were collected May 21. 

Records. UNITED STATES: Colorado: Palisade, May, 3 (holotype and 
paratypes). Utah: Dinosaur National Monument, May, 2. Wyoming: 
State label only, 1. 

Remarks. One of the paratypes has a pair of small, blunt tubercles on 
each side of the pronotum just before the basal impressed line. 

The presence of only a single spur on each fore tibia in the male dis- 
tinguishes nigrocyanea from all other North American species of Lytta 
except moerens. The evident affinity of nigrocyanea with the species of 
the Cyanipennis Subgroup has been mentioned above. 

The type of nigrocyanea is in the collection of the California Academy 



204 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

of Sciences (no. 2601, examined). The paratypes examined are in the 
collections of the Brigham Young University and the California Academy 
of Sciences. According to the original description of nigrocyanea, there 
should be additional paratypes in the collection of the Colorado Agricul- 
tural and Mechanical College at Fort Collins. I have written twice to 
this institution requesting information about these, but neither of my 
letters has been answered. 

Cyanipennis Subgroup 

Bright metallic green, blue, purple, or violaceous. Frontal spot one- 
sixth or less as wide as frontal area between eyes. Head wider than long, 
strongly triangular; tempora prominent; dorsal margin nearly straight; 
vertex somewhat tumid; surface smooth, shiny, moderately densely to 
sparsely punctate; pubescence long, erect. Male antennae with segments 
IV-VII incrassate, subequal in length, bulged on lateral side, triangular; 
VI and VII obviously thicker than rest; VIII-X progressively shorter and 
narrower; VIII similar in shape to VII, about as wide as V. Pronotum 
hexagonal, wider than long; lateral angles well marked; disk flattened 
or impressed before middle, usually foveate on each side at middle; 
base strongly impressed along margin, which is reflexed, the impression 
broadened medianly; surface more finely punctate than on head; punc- 
tures usually denser behind middle than before; pubescence as on head. 
Elytra impunctate; pubescence sparse; female with pubescence shorter 
and sparser than in male and with long setae never present basally. Apex 
of fore and middle tibiae acutely produced on posterior side in male; apex 
of fore-tibiae flared in this position in female. Male fore tibiae each with 
two spins; anterior spur long, heavy, distorted, excavate at base. Male 
posterior middle tibial spur sticklike, somewhat contorted. Outer hind 
tibial spur at most twice as wide as inner spur; truncature oval; inner spur 
flattened. First segment of male fore tarsi strongly modified, bent in 
dorsal view, with a deep, socket-like excavation anteroventrally at base; 
first segment of female fore tarsi strongly cultriform, straight, not exca- 
vate. Tarsal claws moderately curved. Emargination of male fifth ab- 
dominal sternum broad, notched at apex. Female fifth abdominal sternum 
shallowly emarginate. Male sixth sternum with mesal margin of lateral 
lobes as well as an elongate, rectangular median area lightly sclerotized; 
median area with two more or less distinct, parallel rows of long setae, 
these rows extending onto lateral lobes of sternum as fringes; long setae 
usually bent at apex. Emargination of female sixth sternum moderately 
deep, circular or oval. Male genitalia as in Figure 345; gonostvli lacking 
mesal hooks, very conspicuously clothed ventrally with long setae; aede- 
agus with a single ventral hook, this hook strongly recurved. 



SYSTEMATICS AND PHYLOGENY 205 

Remarks. This subgroup contains three very similar forms: nuttalli, 
cyanipennis, and viridana. The last two of these are especially close 
morphologically and, in view of their allopatric distribution, may be 
conspecific. 

The range of the Cyanipennis Subgroup encompasses the main moun- 
tain masses of western North America as far north as the Peace River 
region of Canada and south to the southern limits of the Rocky Moun- 
tains, in New Mexico and Arizona. The range of nuttalli, which includes 
the greater part of the combined ranges of cyanipennis and viridana, is; 
one of the largest in the genus. Its range is also notable in that it extends 
farther east than that of any other North American species of Lytta ex- 
clusive of the species of the subgenus Pomphopoea. 

Key to Species 

1. Elytra generally violet, at least on margins, rarely entirely green; 
male fifth abdominal sternum with lateral lobes angulate (Fig. 219) 

nuttalli (p. 205) 

Elytra lacking violet color; male fifth abdominal sternum with lateral 
lobes rounded 2 

2. Male antennal segments IV-VIII incrassate but not otherwise modi- 
fied; hind trochanters spined or angulate on ventral margin in male, 
angulate or smooth in female; individual setae on under surface of 

thorax entirely piceous cyanipennis (p. 209) 

Male antennal segments IV-VIII flattened and clothed with long, 
erect setae on lateral surface; hind trochanters not spined or angulate; 
individual setae on under surface of thorax colorless apically, piceous 
basally viridana ( p. 213 ) 

Lytta (Poreospasta) nuttalli Say 

Lytta nuttallii Say, 1824, Jour. Acad. Nat. Sci. Philadelphia, vol. 3, p. 300. 

Lytta nuttalli, 4 Say, 1824, American entomology, vol. 1, p. 5, pi. 3, fig. 1. 
LeConte, 1853, Proc! Acad. Nat. Sci. Philadelphia, vol. 6, p. 334. Horn, 1872, 
Rept. U.S. Geol. Surv., 1871, p. 390. Cockerell and Harris, 1925, Proc. Biol. 
Soc. Washington, vol. 38, p. 30, fig. 9. Carruth, 1931, Ent. News, vol. 42, 
p. 54. Knowlton, 1930, Florida Ent., vol. 14, p. 53; 1939, Utah Agr. Exp. Sta. 
Mimeo. Ser. 200, pt. 3, p. 5. Fox, 1943, Canadian Ent., vol. 75, p. 206. 
Peterson, 1951, Larvae of insects, vol. 2, p. 198, fig. C57, A. MacSwain, 1956, 
Univ. California Publ. Ent., vol. 12, p. 87. 

Cantharis fulgifer LeConte, 1847, Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 
vol. 1, p. 90. 

Lytta fulgifer, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia vol. 6, 
p. 334. 

Cantharis nuttalli, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 106; 

4 The specific name nuttalli has been frequently misspelled as nuttali or nutalli, 
but it does not seem necessary to segregate these variants in listing the synonymy. 



206 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

1873, Rept. U.S. Geol. Surv., 1872, 1873, p. 717. Saunders, 1876, Canadian 
Ent., vol. 8, p. 224, fig. 11. Wickham, 1896, Canadian Ent., vol. 28, p. 34. 
Chittenden, 1899, Yearbook U.S. Dept. Agr., 1898, p. 250. Fall, 1901, Trans. 
Amer. Ent. Soc., vol. 27, p. 297. Wickham, 1901, Psyche, vol. 9, p. 151. 
Chittenden, 1903, U.S. Dept. Agr. Div. Ent. Bull. 40, p. 116; 1903, U.S. Dept. 
Agr. Div. Ent. Bull. 43, p. 27, fig. 22. Fall and Cockerell, 1907, Trans. Amer. 
Ent. Soc, vol. 33, p. 210. Gibson, 1912, 42nd Ann. Rept. Ent. Soc. Ontario, 
p. 2, fig. 4. 

Lytta nuttalli var. fulgifera, Ulke, 1875, in Report upon the geographical 
and geological explorations and surveys west of the 100th meridian, vol. 5, 
p. 825. 

Cantharis nuttali [sic] var. fulgifera, Putnam, 1876, Proc. Davenport Acad. 
Nat. Sci., vol. 1, p. 91. 

Cantharis fulgifera, LeConte, 1878, Bull. U.S. Geol. Surv., vol. 4, pt. 2, 
p. 472. 

Head and pronotum brassy green, usually edged with purple, varying 
rarely to brassy violet. Elytra somewhat iridescent, usually metallic 
dark violet, becoming lighter and distinctly brassy toward base and mar- 
gins, the immediate base and margins brassy green; elytra varying occa- 
sionally through brassy violet to brassy green, in which case the margins 
and base are almost always violet. Under surface metallic dark green 
or blue-green, usually grading to purple along edges of sclerites. Femora 
and tibiae metallic green, blue, or purple. Wings nearly colorless. Pubes- 
cence on under surface of thorax distinctly pale, silvery, the individual 
setae entirely colorless or colorless with base piceous; rest of pubescence 
piceous. Length: 7 mm. -21 mm. 

Pronotum (Fig. 189) with lateral angles generally a little more prom- 
inent and acute than in cyanipennis and viridana. Elytra sparsely clothed 
with short pubescence, with some long setae usually present at immediate 
base. 

Male. Antennae as in Figure 112; intermediate segments incrassate 
but not otherwise modified. Hind trochanters (Fig. 163) armed with a 
well-developed spine on ventral margin just basad of middle. Fifth ab- 
dominal sternum (Fig. 219) with lateral lobes angulate. Sixth sternum 
as in Figure 268. Genitalia as in Figure 345. 

Female. Antennae as in Figure 150. Hind trochanters angulate on ven- 
tral margin near middle. Sixth abdominal sternum as in Figure 285. 

Type locality. Of nuttalli, Missouri Territory; of fulgifera, near Longs 
Peak, Colorado. 

Geographic distribution. Central Alberta and northern Minnesota south 
to California, Arizona, and New Mexico. ( See Fig. 54. ) Limited to higher 
elevations in the southern half of its range. The Sierra Nevada popula- 
tion appears to be disjunct from the main range of the species. 

Saunders' (1876) report that nuttalli is "extremely abundant" in Kansas 



SYSTEMATICS AND PHYLOGENY 



207 



is evidently erroneous as I have seen no specimens from that state and 
it is unlikely that the species occurs so far south east of the Rocky Moun- 
tains. There is a specimen in the Chicago Natural History Museum 
labeled Toronto, Ontario (September 4, 1930, F. H. Daniels), which I 
have disregarded pending verification in the form of additional material 
from that locality. 

Seasonal distribution. May 31 — September 21. Collected most fre- 
quently in June and July. As in the other two species of the subgroup, the 
seasonal distribution does not seem to be affected by latitude. 

Records. CANADA: Alberta: Cardston, 1; Chin, June, 1; Edmonton, 
3 (2 in July); Iron Springs, June, 1; Laggen, June, 1; Lethbridge, June, 
2; Medicine Hat, June, July, 16; Scandia, June, 57; Stavely, July, 1. 
Manitoba: Aweme, June, July, 8; McCreary, June, 5; Melita, July, 1; 
Sandilands, September, 1; Winnipeg, 3. Saskatchewan: Attons Lake, Cut 
Knife, June, 3; Indian Head, July, 2; Lloydminster, June, July, 4; Moose 
Jaw, 1; North Portal, 3; Oxbow, June, 3; Penzance, 2; Roche Percee, July, 
2; Saskatoon, June, 5; Weyburn, June, 2. 




Fig. 54. Distribution of Lijtta nuttalli. 



208 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

UNITED STATES: Arizona: Kaibab National Forest, June, 1; Spring- 
erville, July, August, 9; White Mountains, 10,000 ft., July, 3. California: 
State label only, 2; Inyo County, July, 1; Monache Mountain, Tulare 
County, July, 4; Sardine Creek, 8500 ft., Mono County, June, 1; Sonora 
Pass, 9000 ft.-l 1,000 ft., Alpine County, June, July, 282; Twin Lake, Mono 
County, July, 1. Colorado: State label only, 45; Antonito, July, 13; Bent's 
Fork (LeConte, 1853); Cochetopa National Forest, September, 1; Colo- 
rado Springs, July, 1; Creede, 1844 ft., August, 4; Denver, 8; Eastonville, 
August, 2; Fort Collins, June, 15; Georgetown, 2; Gilpin County, 9500 
ft., July, 2; Great Sand Dunes, Alamosa County, July, 1; Larkspur, July, 
10; near Leadville, August, 6; near Longs Peak (LeConte, 1847); North 
Park, June, 4; Palmer Lake, June, 2; Pingree Park, August, 3; Pinnacle, 1; 
San Isabel National Forest, August, 3; South Park (Ulke, 1875); Welling- 
ton, July, 1; Willow Park (not located), 3. Idaho: Atlanta, 7800 ft. 
(LeConte, 1878); Snake River (Horn, 1873, p. 717). Minnesota: State 
label only, 2; Itasca [State] Park, 12 (8 in June, July); St. Anthony 
Park, August, 1. Montana: State label only, 20; Bozeman, August, 10; 
Carlyle, July, 1; Havre, June, 1; Helena, July, 2; Lewis and Clark County, 
July, 2; Lima, June, 1; Moccasin, July, 1; Rock Creek Lake, Powell 
County, July, 2; Sheridan, June, July, 6. Nebraska: (Chittenden, 1903, 
p. 27). New Mexico: State label only, 9; Beulah, August, 2; near Hot 
Springs, Las Vegas, 7000 ft., July, 1. North Dakota: State label only, 1; 
Custer, 2; Fessenden, June, 5; Forman, July, 1; Glenburn, July, 1; Hank- 
inson, August, 2; Mandan, June, 6. South Dakota: State label only, 1; 
Black Hills, 1; Brookings, 13 (1 in July); Buffalo (Carruth, 1931); Custer, 
July, 2; Faulkton, Leola, Newell, Selby, and Sisseton Indian Reservation 
(Carruth, 1931); Volga, 1. Utah: State label only, 2; Bear Lake, 1; Du- 
chesne, July, 1; Fish Lake, June, July, 28; Fruitland, July, 1; Lambs 
Canyon, Salt Lake County, August, 14; Logan, July, 1; Lost Lake, August, 
2; Mill Creek Canyon, Salt Lake County, July, 2; Payson Canyon summit, 
1; Timothy Creek, 7500 ft., 10 mi. north of Altonah, Duchesne County, 
June, 1; Valley View, Mount Nebo, Juab County, July, 1. Wyoming: State 
label only, 2; Bamforth Lakes, 10 mi. northwest of Laramie, July, 13; 
Big Horn Mountains, July, 1; Evanston, June, 1; Grand Teton National 
Park, July, August, 7; Hulett, July, 2; Laramie, 2; Wind River Basin ( Put- 
nam, 1876); Yellowstone National Park, 27 (21 in July). 

Remarks. The violet color of the elytra will distinguish nuttalli from 
cyanipennis and viridana in almost all cases. In the absence of this color, 
males of nuttalli can be distinguished from those of cyanipennis by the 
form of the fifth abdominal sternum and from those of viridana by this 
same character as well as by the absence of long pubescence on the 
antennae. Atypically colored females of nuttalli are best separated from 
specimens of cyanipennis by the pale pubescence of the under surface 



SYSTEMATICS AND PHYLOGENY 209 

of the thorax and from specimens of viridana by the distinctly angulate 
hind trochanters. 

Aside from the variation in elytral color, which is not geographic, 
nuttalli is remarkably stable throughout its range. No structural variation 
worthy of note has been observed other than that involving two speci- 
mens from Edmonton, Alberta, which appear to be hybrids between 
nuttalli and viridana (see discussion of viridana). 

The first instar larva of nuttalli has been described by MacSwain 
(1956). The type has been lost. As neotype I have designated a male 
from Brookings, South Dakota, in the collection of Cornell University. 

Bionomics. Except for reports of occasional injury to oats and barley 
in western Canada ( Gibson, 1912 ) and North Dakota ( Chittenden, 1903, 
p. 116), nuttalli has been recorded only from species of Leguminosae. 
Among the records which have come to my attention are the following: 
Astragalus in western Canada (Gibson, 1912) and California, Caragana 
arborescens in Utah, Caragana sp. in North Dakota and Saskatchewan 
(Penzance), Cnemidophacos pectinatus and Diholcos bisulcatus in Sas- 
katchewan (Fox, 1943), Thermopsis montanus in Utah, and Vicia ameri- 
cana in western Canada (Gibson, 1912). There have also been some 
reports of feeding on cultivated beans and beets ( Chittenden, 1899; 1903, 
p. 116; 1903, p. 27; and Gibson, 1912). 

Fox (1943) has reported that adults of nuttalli feeding on Cnemido- 
phacos and Diholcos in Saskatchewan are attacked by the mirid bug 
Hadronema militaris Uhler. 

Lytta (Poreospasta) cyanipennis (LeConte) 

CantJiaris cyanipennis LeConte, 1851, Ann. Lye. Nat. Hist. New York, vol. 
5, p. 160. Horn, 1873, Proc. Amer. Philos. Soc.,' vol. 13, p. 107; 1873, Rept. 
U.S. Geol. Surv., 1872, 1873, p. 717. LeConte, 1878, Bull. U.S. Geol. Surv., 
vol. 4, p. 472. Wickham, 1896, Canadian Ent., vol. 28, p. 34. Fall, 1901, Occas. 
Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc., 
vol. 27, p. 297. Wickham, 1902, Bull. Lab. Nat. Hist. State Univ. Iowa, vol. 
5, p. 301. Gibson, 1912, 42nd Ann. Rept. Ent. Soc. Ontario, p. 4. 

Lytta salicis LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 333; 1866, Smithsonian Misc. Coll., vol. 6, no. 167, p. 162. 

Lytta cyanipennis, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, 
vol. 6, p. 333. Horn, 1872, Rept. U.S. Geol. Surv., 1871, p. 390. Knowlton, 
1930, Florida Ent., vol. 14, p. 53; 1939, Utah Agr. Exp. Sta. Mimeo. Ser. 200, 
pt. 3, p. 5. Knowlton and Taylor, 1952, Utah Agr. Exp. Sta. Mimeo. Ser. 389, 
p. 15. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 86. Hicks, 1959, 
Check-list and bibliography on the occurrence of insects in birds' nests, p. 72. 

Differs from nuttalli as follows. 

Elytra metallic dark purple, blue, or green, never violet. Brassy, iri- 
descent quality almost always lacking. Under surface the same color as 



210 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 



elytra or the same color with an admixture of green, rarely at all brassy. 
Pubescence entirely dark. Length: 8 mm. -20 mm. 

Male. Fifth abdominal sternum (Fig. 220) with lateral lobes broadly 
rounded, not angulate. Elytra frequently with long, erect setae on basal 
half to two-thirds. Hind trochanters with a well-developed spine on ven- 
tral margin at or basad of middle ( Fig. 164 ) or with spine reduced to a 
tooth or acute angulation ( rarely obsolete ) and located distad of middle 
of ventral margin (Fig. 165). 

Female. Hind trochanters with ventral margin varying from distinctly 
angulate (Fig. 166) to smoothly rounded (Fig. 167). 

Type locality. Of cyanipennis, Oregon; of salicis, Salt Lake City, Utah. 

Geographic distribution. Southern British Columbia south to northern 
California, southern Utah, and southern Wyoming. ( See Fig. 55. ) Limited 
to higher elevations in the southern part of its range. 

Seasonal distribution. March 24 — August 18. 




Fig. 55. Distribution of Lytta cyanipennis (circles) and Lytta viridana (squares). 



SYSTEMATICS AND PHYLOGENY 211 

Records. CANADA: British Columbia: Clinton, June, 1; Crown Lake, 
June, 8; Douglas Lake, June, 5; Ducks (Gibson, 1912); Hulcar, May, 2; 
Lac du Bois, Kamloops, 3700 ft., July, 3; Lytton, June, 1; Midday Valley, 
June, July, 12; Okanagan Landing, May, 1; Oliver, June, 14; Pavilion 
Lake, June, 14; Penticton, June, 1; Princeton, May, 1; Similkameen, May, 
3; Seton Lake, June, 1; Six Mile Creek, May, 3; Spious Creek (not lo- 
cated), June, 1; Summerland, June, 1; Vernon, May, 8; Westbank, June, 1. 

UNITED STATES: California: Bucks Lake, Plumas County, June, 
July, 5; Dunsmuir, 1; MacDoel, May, 1; Middle Fork, American River, 
Placer County, August, 1; Sierra Nevada, 2. Idaho: Atlanta, 4; Avon, 
June, 1; Bannock Pass, Lemhi County, July, 23; Bloomington Lake, 
Wasatch Mountains, 8200 ft., July, 4; Brundage Lookout, Payette Na- 
tional Forest, August, 1; Caldwell, 2375 ft., May, 1; Deary, 2775 ft., May, 
25; Fort Sherman, June. 1; Granite, July, 4; Joel, May, 3; Lakefork, Valley 
County, July, 2; Lewiston, 5000 ft., March-May, 5; McCall, June, 1; 
Moscow, May, June, 9; Moscow Mountain, 3000 ft., May, 1; St. Maries, 
June, 1; Sandpoint, 2086 ft., June, 4; Stanley, June, 1; Teton, June, 1; 
Warm Lake, June, 3; Winchester, June, 1. Montana: Beaverhead County, 
May, 1; Bitter Root Valley, 1; Bozeman, June, 2; Butte, 1; Gallatin County, 
June, July, 8; Glacier National Park, June, July, 9; Madison County, July, 
11; Missoula, 5; Rock Creek Lake, Powell County, July, 1; 5 mi. east of 
Saint Regis, June, 16; Thompson Falls, June, 1. Nevada: Elko, June, 3. 
Oregon: Arlington, May, 2; Burns, June, 2; Durkee, June, 4; Enterprise, 
July, 1; Fish Lake, Steens Mountains, 7000 ft., July, 1; Klamath Falls and 
17 mi. east of Klamath Falls, June, July, 2; Langden Lake, Umatilla 
County, June, 1; Lexington, May, 1; Milton, 1500 ft., May, 1; Moro, April, 
7; Ochoco National Forest, June, 2; Princeville, June, 1; Silver Creek, July, 
6. Utah: Alta, Little Cottonwood Canyon, 9000 ft.-10,000 ft., July, August, 
200; Aspen Grove, Mount Timpanogas, July, 13; Bell Canyon, Salt Lake 
County, July, 1; Brighton, 9000 ft.-10,000 ft., July, 45; Cedar Breaks, July, 
4; Chalk Creek, Summit County, July, 1; Fish Lake, 8600 ft., June, July, 
48; Logan Canyon, June, August, 17; Mill Creek Canyon, Salt Lake 
County, July, 5; Mueller Park, Davis County, June, 1; Provo and Provo 
Canyon, May, July, 19; Spanish Fork, 1; Strawberry Valley, June, 1; Tony 
Grove Ranger Station, Cache County, July, 1; west of Woodruff Moun- 
tains, 8000 ft., Rich County, June, 1. Washington: Alderdale, May, 1; 
Dayton, May, 4; Dryden, May, 3; Easton, 9; Kiona, May, 2; Mount Rainier 
National Forest, June, 1; Northport, June, 6; Pullman, 3; Tieton, May, 2; 
Walla Walla, May, 28; Yakima County, July, 2. 

Remarks. The length of the elytral pubescence is extremely variable 
in the male. In the southern half of the range (except in California) the 
elytral pubescence is short throughout, or if long setae are present, they 
are restricted to the immediate base of the elytra. In a few specimens 



212 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

from this area long setae are present as far apicad as the basal fourth of 
the elytra. In northern Idaho about two-thirds, and elsewhere within the 
northern half of the range (and in California) about half, of the speci- 
mens have distinctly hairy elytra, with long setae along the sutural third 
of each elytron from the base to the middle or apical third. Other speci- 
mens from the northern area have entirely short pubescence, and all 
degrees of intermediacy between the extremes is represented in still 
others. 

Throughout the greater part of the range of cyanipennis the male hind 
trochanters have a well-developed spine located at or basad of the 
middle of the ventral margin (Fig. 164). In Utah, southeastern Idaho, 
and southern Wyoming it is reduced to a short tooth (Fig. 165) located 
distad of the middle of the ventral margin or (rarely) is absent. Modi- 
fication of the female hind trochanters is correlated geographically with 
that of the male: where the male trochanters have a well-developed 
spine, the ventral margin of the female trochanters is distinctly angulate; 
where the male spine is reduced, the ventral margin of the female tro- 
chanters is rounded. Series of specimens from Nevada (Elko), south- 
western Idaho (Atlanta and Stanley), northwestern Wyoming (Yellow- 
stone and Grand Teton national parks), and the Gallatin Mountain 
region of Montana show, to a greater or lesser extent, intermediate de- 
velopment of the trochanteral modification. 

The first instar larva of this species has been described by MacSwain 
( 1956 ) . The type of cyanipennis, as well as that of solids, is in the Mu- 
seum of Comparative Zoology. 

Bionomics. The type material of the synonym salicis was reportedly 
taken on willow, but this record has never been verified and is probably 
wrong. All other food plants recorded for cyanipennis belong to the family 
Leguminosae. One specimen examined from Idaho was collected on 
Vicia, and there are several records of Lupinus from British Columbia, 
Idaho, Montana, and Utah. In addition, Gibson (1912) reported injury 
to peavine [Lathyrus ?], beans, and peas in British Columbia. 

In the Wasatch Range of Utah I have frequently found cyanipennis 
feeding on a species of Lupinus growing at elevations between 8000 ft. 
and 10,000 ft. The beetles feed on the petals of the flowers in preference 
to the leaves, although they readily accept leaves if flowers are not 
available. In captivity they feed more or less continuously, day and night. 
Their fecal material, which is emitted in large quantity, contains great 
amounts of undigested plant tissue; it is presumably this inefficient utili- 
zation of food that accounts for the unusually ravenous habits of the 
beetles. Efforts to maintain the beetles on plants other than the species 
of Lupinus upon which they feed in nature were uniformly unsuccessful, 
although in one instance some individuals fed sparingly on the leaves of 



SYSTEMATICS AND PHYLOGENY 213 

a different species of Lupinus and on the leaves of fresh lettuce. Alfalfa 
and several species of Compositae were offered on various occasions and 
were always refused. 

Hicks (1959), citing the unpublished doctoral dissertation of W. L. 
Jellison, records cyanipennis from the nest of the magpie Pica pica hud- 
sonica. 

Lytta (Poreospasta) viridana LeConte 

Lytta viridana LeConte, 1866, Smithsonian Misc. Coll., vol. 6, no. 167, 
p. 162. Horn, 1872, Rept. U.S. Geol. Surv., 1871, p. 390. Fox, 1943, Canadian 
Ent., vol. 75, p. 206. 

Cantharis viridana, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 107. 
LeConte, 1879, Bull. U.S. Geol. Surv., vol. 5, pt. 3, p. 506. Fall, 1901, Trans. 
Amer. Ent. Soc, vol. 27, p. 297. Wickham, 1902, Bull. Lab. Nat. Hist. State 
Univ. Iowa, vol. 5, p. 301. Fall and Cockerell, 1907, Trans. Amer. Ent. Soc., 
vol. 33, p. 210. Gibson, 1912, 42nd Ann. Rept. Ent. Soc. Ontario, p. 15. 

Differs from cyanipennis as follows. 

Individual setae on under surface of thorax colorless apically, piceous 
basally (entirely piceous in cyanipennis). 

Male. Antennal segments IV-VIII flat and conspicuously clothed with 
long, erect setae on outer surface. Hind trochanters almost invariably 
smooth on ventral margin, without an indication of a spine. Elytra with 
pubescence usually slightly longer at base than in female but only very 
rarely with any long, erect setae. 

Female. Hind trochanters with ventral margin never angulate. 

Type locality. "Rocky Mountains, from the Black Hills northward into 
the Hudson Bay Territory" (LeConte, 1866). 

Geographic distribution. Peace River region of British Columbia and 
southern Manitoba south through eastern Wyoming, Colorado, and south- 
eastern Utah to northern Arizona and New Mexico. ( See Fig. 55. ) Limited 
to higher elevations in the southern part of its range. Consistently allo- 
patric with cyanipennis. 

I have seen a few specimens supposedly from Kansas, but I doubt that 
they are correctly labeled. Tanner and Hayward's ( 1934 ) record of 
viridana from Utah was based on misidentified specimens of Linsleya 
sphaericollis ( Say ) . 

Seasonal distribution. May 20 — August 14. 

Records. CANADA: Alberta: Bilby, June, 1; Calgary, May, 25; Coutts, 
June, 2; Edmonton, June, July, 14 (including 2 viridana x nuttalli hy- 
brids); Laggan, May, 1; Medicine Hat, May, June, 38. British Columbia: 
Fort St. John, June, 1; Taylor, June, 20. Manitoba: Aweme, June, 2. 
Saskatchewan: Asquith, May, 1; Christopher Lake, May-July, 21; Ogema, 
June, 1; Oxbow, June, 3; Penzance, June, 2; Prince Albert National 



214 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Park, June, 3; Rivercourse Post Office, May, 2; Rudy (Gibson, 1912); 
Saskatoon, June, 1. 

UNITED STATES: Arizona: San Francisco Mountains, June, July, 20 
(9 at Snow Bowl). Colorado: State label only, 170; Creede, 8844 ft., 
August, 34; Dumont, June, 1; Estes Park, July, 1; Fort Collins (Wickham, 
1902); Georgetown, 8300 ft.-8600 ft., June, 6; Grand Lake, August, 4; 
Green Mountain Falls, August, 1; Gunnison, 7500 ft., June, 2; Jefferson, 
July, 5; 18 mi. north of Leadville, June, 3; Little Beaver (Wickham, 1902); 
southwest of Montrose ( Wickham, 1902 ) ; near Nederland, June, 8; North 
Park, June, 1; Ohio, July, 2; Ouray (Wickham, 1902); Palmer Lake, June, 
7; Ragged Mountain, Gunnison County, June, 1; Rico (Wickham, 1902); 
Rio Grande, 10 mi. southwest of Creede, July, 1; Rocky Mountain Na- 
tional Park (Fall River Entrance and Hidden Valley), June, 4; Russel, 
June, 1; Sheep Lake, Rocky Mountain National Forest, June, 1; [La] 
Veta Pass, 1; Ward, July, 6; West Cliff (Wickham, 1902); Willow Park 
(not located), 15. New Mexico: Abiquiu, 1; Harvey s Ranch, Las Vegas 
Range (Fall and Cockerell, 1907); Jemez Mountains, May, 11. Utah: 
Miners Peak [Mountain], Wayne County, July, 3. Wyoming: State label 
only, July 1; 25 mi. west of Cheyenne, June, 1; Laramie, July, 2. 

Remarks. The only consistent basis for separation of females of viridana 
and cyanipennis is the difference in the color of the pubescence of the 
under surface of the thorax. This difference is best observed when 
the pubescence is viewed in profile against back lighting. Males of the 
two species are easily separated by use of the antennal and trochanteral 
characters indicated in the key. The character of elytral pubescence is 
of no real value in making determinations because of the great variability 
in cyanipennis. An occasional green female of nuttalli may be confused 
with viridana but can be distinguished by its definitely angulate hind 
trochanters. 

In view of the great similarity between viridana and cyanipennis, the 
allopatry of the two forms suggests that they may represent geographical 
races of the same species. Such a hypothesis is supported by the fact 
that the population of cyanipennis in southern Idaho, Wyoming, and Utah 
approaches viridana with respect to reduction of the modification of the 
hind trochanters. However, there is no evidence of intergradation between 
viridana and cyanipennis in characters of the male antennae and color 
of the thoracic pubescence. The problem of the relationship of the two 
forms requires a great deal of additional study, pending which it seems 
most reasonable to consider them as separate species. This decision is 
in line with the knowledge that nuttalli, which is sympatric with both 
viridana and cyanipennis and is unquestionably specifically distinct from 
both, shows only slight structural differentiation from either of them. 



SYSTEMATICS AND PHYLOGENY 215 

Hybridization of viridana and nuttalli is indicated by two males from 
Edmonton, Alberta, collected July 12, 1917, by F. S. Carr. These speci- 
mens have the violet elytra of nuttalli and the unmodified hind trochanters 
and rounded lateral lobes of the fifth abdominal sternum of viridana. 
One has the antennae of nuttalli, while the other has antennae like 
those of viridana but with the intermediate segments less flattened 
laterally and with the erect setae shorter and sparser than usual. In 
addition to these specimens, I have seen from Edmonton 12 specimens of 
typical viridana (including 1 specimen with collection data identical 
to those of the hybrids ) and 3 specimens of typical nuttalli. Inasmuch as 
the two species occur sympatrically over a large area and apparently do 
not have food plant differences, the occurrence of occasional hybrids is 
not particularly surprising. 

Bionomics. Apparently restricted to Leguminosae. Fox (1943) has re- 
ported that in Saskatchewan viridana occurs with nuttalli on Cnemido- 
phacos pectinatus and Diholcos bisulcatus. Other food plants are Cara- 
gana at Penzance (again with nuttalli) and peavine [Lathyrus ?] at 
Prince Albert National Park, both localities in Saskatchewan. 

Childi Subgroup 

Black, without metallic luster. Frontal spot diamond-shaped. Pronotum 
frequently in part orange. Head quadrate, as long as wide; surface sub- 
glabrous, irregularly punctate, with some very fine punctures intermixed 
with larger ones. Male antennae with segment I densely punctulate on 
ventral side; intermediate segments incrassate or not. Pronotum hex- 
agonal (molesta) or subquadrate (childi); surface more irregularly, 
sparsely punctate than head, subglabrous. Scutellum rounded. Elytra im- 
punctate, essentially glabrous. Outer hind tibial spur cylindrical or nearly 
so, four times as wide as inner spur; truncature broadly oval. Emargina- 
tion of male fifth abdominal sternum broadly triangular. Male genitalia 
with gonostyli lacking mesal hooks, clothed ventrally with long setae; 
aedeagus with a single ventral hook or none. 

Remarks. This subgroup contains the species molesta and childi. Its 
range extends from central California south to the Cape Region of the 
peninsula of Baja California. 

Key to Species 

1. Pronotum hexagonal (Fig. 190); male antennae (Fig. 113) clavate; 
male hind trochanters each armed with a ventral spine near apex; 
female sixth abdominal sternum (Fig. 286) with a broadly rounded 
emargination molesta ( p. 216 ) 



216 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF LyttCl 

Pronotum subquadrate (Fig. 191); male antennae not clavate, the 
intermediate segments incrassate; male hind trochanters not spined; 
female sixth abdominal sternum (Fig. 287) with a narrow, U-shaped 
emargination childi ( p. 218 ) 

Lytta (Poreospasta) molesta (Horn) 

Cantharis molesta Horn, 1885, Trans. Amer. Ent. Soc, vol. 12, p. 111. Fall, 
1901, Trans. Amer. Ent. Soc, vol. 27, p. 298. 

Lytta molesta, Van Dyke, 1929, Bull. Brooklyn Ent. Soc, vol. 24, p. 131. 
MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 89, pi. 15. 

Black. Frontal spot nearly always more than half as wide as frontal 
area between eyes, rarely half as wide. Pronotum orange or reddish 
orange with anterior margin of disk broadly black and with a large 
black discal blotch extending from basal margin to before middle, as in 
Figure 190, or with orange arch so formed narrowly or broadly broken 
medianly; very rarely with pronotum entirely black. Wings light brown 
with area basad of 1A nearly colorless. Pubescence silvery on under 
surface of thorax, piceous elsewhere. Length: 11 mm.-22 mm. 

Head with sides a little more divergent above eyes than in childi and 
with tempora therefore more prominent; surface smooth, sparsely punc- 
tate. Pronotum (Fig. 190) hexagonal, one-fifth to one-fourth wider than 
long; basal margin more strongly emarginate medianly than in childi; 
surface smooth or with some fine alutaceous markings, especially basally. 
Tarsal claws moderately curved. 

Male. Antennae (Fig. 113) reaching one segment beyond base of pro- 
notum; form nearly as in female, the intermediate segments not incras- 
sate; segments III-VIII subequal in length; III as wide as VI, IV and V 
a little narrower; VI-X becoming progressively slightly wider; IX and X 
a little shorter than VIII; XI at most four-fifths as long as IX and X 
combined. Hind trochanters each armed with a spine on ventral margin 
near apex; ventral margin nearly straight; apex of trochanter truncate. 
Posterior margin of middle tibiae produced as a broad lamella reaching 
beyond middle of posterior tibial spur. Posterior middle tibial spur heavy, 
contorted, projecting at a wide angle, truncate at apex. Sixth abdominal 
sternum as in Figure 269. Genitalia as in Figure 346; aedeagus with 
ventral hook well developed. 

Female. Antennae (Fig. 151) reaching about middle of pronotum; 
segments III-X becoming progressively slightly wider. Sixth abdominal 
sternum (Fig. 286) with a broadly rounded emargination. Genitalia with 
gonostyli very short. 

Type locality. "California . . . probably near Visalia" (Horn, 1885). 



SYSTEMATICS AND PHYLOGENY 



217 



Geographic distribution. Central Valley of California from Contra 
Costa County to Kern and Tulare counties. ( See Fig. 56. ) Recorded from 
Washington by Fall (1901, p. 298) but undoubtedly in error. 

Seasonal distribution. April 3 — July 1. 

Records. UNITED STATES: California: State label only, 11; Auberry, 
April, 1; 10 mi. north of Blackwells Corner, Kern County, April, 64; Brent- 
wood and 3 mi. southwest of Brentwood, April, 18; El Mirador, Tulare 
County, April, 2; Fresno, June, July, 6; Kern County, 7; Kings County, 
June, 5; Lanes Bridge, 5 mi. north of Fresno, April, 3; Madera, April, 1; 
6 mi. north of Merced, April, 3; Panoche Canyon, Fresno County, April, 
1; Springville, April, 2; 14 mi. north of Wasco, April, 52. 

Remarks. The pronotum is entirely black in one of the specimens from 
6 mi. north of Merced. In all other specimens of the species that I have 
seen it is extensively marked with orange. 

The first instar larva of molesta has been described by MacSwain 
( 1956 ) . The type is in the collection of the Academy of Natural Sciences 
of Philadelphia. 

Bionomics. There are records from Lupinus ( Leguminosae ) in Contra 
Costa, Kern, and Fresno counties. The specimen from Auberry, Fresno 
County, was collected on filaree [Er odium cicutarium ( Geraneaceae ) ] . 




Fig. 56 (left). Distribution of Lytta molesta. 
Fig. 57 (right). Distribution of Lytta child*. 



218 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

Lytta (Poreospasta) childi LeConte 

Lytta childii LeConte, 1857, in Reports of explorations and surveys . . 
from the Mississippi River to the Pacific Ocean, vol. 12, book 2, pt. 3, p. 52 

Cantharis childii, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 108 
1894, Proc. California Acad. Sci., ser. 2, vol. 4, p. 356. Fall, 1901, Occas 
Papers California Acad. Sci., vol. 8, p. 185; 1901, Trans. Amer. Ent. Soc, vol 
27, p. 297. 

Lytta maculicollis Van Dyke, 1929, Bull. Brooklyn Ent. Soc, vol. 24, p. 130. 
Not Klug, 1825, Verh. Kaiserlichen Leopoldinsch-Carolinischen Akad. Natur- 
forsch., vol. 12, p. 445. New sijnonymy. 

Lytta childi, MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 89. 

Black. Frontal spot varying from less than one-third to nearly half as 
wide as frontal area between eyes. Pronotum varying from entirely black 
to largely reddish orange ( Fig. 58 ) . Antennae, prosternum, and tarsi often 
reddish when pronotum is extensively marked with orange. Metasternum 
with midline rarely reddish. Wings light brown, with area basad of 1A 
nearly colorless. Pubescence piceous. Length: 10 mm.-20 mm. 

Head surface sometimes weakly alutaceous or canaliculate, especially 
frontal area, sparsely to rather densely punctate. Pronotum (Fig. 191) 
subquadrate, varying from slightly wider than long to slightly longer 
than wide; surface varying from almost smooth to strongly, transversely 
alutaceous. Tarsal claws weakly curved. 

Male. Antennae reaching one to three segments beyond base of 
pronotum; segments III— VII incrassate, bulged on lateral side; IV and 
V largest; VIII-X narrower than preceding segments, becoming progres- 
sively shorter; XI at least as long as IX and X combined, longer than 
first segment of middle tarsus. Hind trochanters not angulate or spined 
on ventral margin, subtruncate at apex. Mesal surface of fore tibiae 
lined with dense, golden pubescence. Middle tibial spurs spiniform, not 
modified. Fifth abdominal sternum as in Figure 221. Sixth sternum as 
in Figure 270; lateral lobes of sternum curving ventrad at apex. Genitalia 
as in Figure 347; aedeagus with ventral hook represented by an apical 
knob. 

Female. Antennae reaching base or basal third of pronotum; segments 
III— VIII subequal in width; IX and X slightly wider. Sixth abdominal 
sternum (Fig. 287) with a narrow, U-shaped emargination medianly. 
Genitalia with gonostyli elongate. 

Type locality. Of childi, San Francisco, California; of maculicollis, 
"near Panoche Creek," Fresno County, California (Van Dyke, 1929). 
The labels of the type series of maculicollis read Panoche Canyon rather 
than Panoche Creek. 

Geographic distribution. Santa Cruz, Mendocino, Sacramento, and 
Tuolumne counties south through the Central Valley to Ventura and Los 



SYSTEMATICS AND PHYLOGENY 219 

Angeles counties, California; thence, based on records of Horn (1894), 
south along the coast to southern Baja California Sur (Isla Santa Mar- 
garita). (See Fig. 57.) 

Seasonal distribution. March 8 — July 7. 

Records. UNITED STATES: California: State label only, 15; Antelope, 
April, 14; 10 mi. north of Blackwells Corner, Kern County, April, 8; Brent- 
wood and 3 mi. southwest of Brentwood, April, 36; By Pass, Yolo County 
(not located), April, 2; Byron, April, 44; Camp Taylor, Marin County, 
June, 1; Coalinga, March, 7; Clovis, July, 1; Davis and north of Davis, 
April-May, 47; 5 mi. south of Democrat Springs, Kern County, April, 11; 
El Mirador, Tulare County, May, 6; Fresno, June, 11; 9 mi. north of 
Fresno, April, 13; 15 mi. north of Fresno, April, 36; Friant, April, 1; 
Jamestown, April, 20; Kern County, March, 1; 11 mi. south of Kettleman 
City, April, 3; La Panza, San Luis Obispo County, May, 1; Lindsay, May, 
2; Los Angeles, 205 ft, March, 1; Los Angeles County, 6 (1 in April); 
Lost Hills, Kern County, April, 1; McClure Valley, Kings County (not 
located), March, 1; Madera, May, 35; Mendocino County (Fall, 1901); 
Mount Tamalpais, Marin County, July, 2; Napa, June, 1; Pacheco Pass, 
Merced County, June, 1; Panoche area (Panoche, Panoche Creek, Pano- 
che Canyon, and Pinoche [Panoche] Hills, Fresno County; Little Pinoche 
[Panoche] Creek, San Benito County), March-April, 76; San Diego 
(Horn, 1894); San Francisco (LeConte, 1857); Santa Cruz, June, 3; 
Simmler, April, 1; Sonoma County, 1; Springville, April, 7; Stanislaus 
[County], April, 2; Taylor State Park, Marin County, May, 1; Tulare, 
April, 2; Ventura, April, 4; Visalia, June, 22; 14 mi. west of Wasco, April, 
6; Woodland, May, 1. 

MEXICO: Baja California Norte: San Julio (Horn, 1894). Baja Cali- 
fornia Sur: Margarita Island [Isla Santa Margarita] (not mapped) 
(Horn, 1894). 

Remarks. The color of the pronotum varies geographically. The varia- 
tion is continuous, but for the purpose of analysis it is convenient to 
recognize a number of arbitrary color classes, to each of which a value 
is assigned reflecting the strength of development and extensiveness of 
the orange marking or markings (Fig. 58). In class 1 the pronotum is 
entirely black. In classes 2 and 3 the lighter areas shown are piceous and 
are either invisible macroscopically or not particularly conspicuous. In 
classes 4-6 they are distinctly orange. By assigning each specimen ex- 
amined to its appropriate color class, an average value has been derived 
for each sample of specimens which serves as an index of "orangeness" 
of the pronotum. These averages are arranged geographically in Figure 
59. The bar graphs in this figure show the frequency distribution of the 
color classes in each sample, the distance between base lines being equal 



220 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Llftta 





4 5 

Fig. 58. Pronotal color classes in Lytta childi. 



to 100 per cent of the sample. The number of specimens in each sample 
is indicated on the graphs. The shaded area on the map is the range of 
molesta. 

As shown in Figure 59, variation in pronotal color in childi has the 
pattern of a centripetal cline for increased development of orange with 
its center in the Central Valley of California. The most interesting aspect 
of this variation is that in the Central Valley childi occurs sympatrically 
with molesta, which has a pronotal color pattern much like that of 
class 5 of childi. Thus childi tends to resemble molesta in color where 
the two species occur together but loses this resemblance where the 
latter species does not occur. 

Similar examples of character convergence have been described in 
other groups of animals. In the butterflies, particularly the heliconids, 
numerous examples are known, most of these apparently accountable 
for on the basis of Mullerian or Batesian mimicry. In a recent review of 
snake mimicry, Hecht and Marien (1956) have described a particularly 
clear-cut case of convergence involving a Lampropeltis mimic of a coral 
snake. Sympatrie convergence of the frogs Rana clamitans and septen- 



SYSTEMATICS AND PHYLOGENY 



221 



trionalis has been described and discussed by Moore ( 1952 ) . In this 
last example the convergence presumably results from the two species 
of frogs being in the same habitat and thus being subject to the same 
selection pressures. 

Available evidence indicates that the beetles childi and molesta also 
share the same habitat. Both species have been collected on Lupinus at 
several points within the area of sympatry. They presumably occur on 
the same species of Lupinus as several mixed series of the two have been 
examined. Moreover, they have the same seasonal distribution. Under 
these circumstances it is not difficult to conceive of one or more of 
the physical factors of the environment producing parallel phenotypic 



I 2 3 4 B 6 



64 


80 


20 

1 i—i 




Fig. 59. Geographic variation in Lytta childi. For explanation see text. 



222 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF hljtta 

or (through natural selection) genotypic responses in the two species. 
The ecological distinctness of the Central Valley from the surrounding 
areas lends some support to this explanation. However, the possibility 
that some form of Miillerian mimicry is operating cannot be overlooked. 
If mimicry is involved, there is presumably a predator exerting a pressure 
on the two species in the Central Valley but not in the peripheral area. 
Batesian mimicry seems to be ruled out as it is very unlikely, in view of 
their close relationship, that one species is more distasteful than the 
other. In addition, it appears that neither of the species occurs more 
abundantly than the other, which is in violation of one of the primary 
conditions under which Batesian mimicry can be operative. Hybridiza- 
tion, another possible explanation of sympatric convergence, does not 
seem to be involved in the present case as there is no evidence of inter- 
gradation between childi and molesta in any of their numerous structural 
differences. 

The first instar larva of childi has been described by MacSwain (1956). 
The type of childi is in the Museum of Comparative Zoology, and the 
type of maculicollis is in the collection of the California Academy of 
Sciences (no. 2602, examined). 

Bionomics. Specimens have been recorded from Lupinus (or "lupine") 
( Leguminosae ) at the following localities in California: Panoche Creek, 
Fresno County (Van Dyke, 1929, as maculicollis); 3 mi. southwest of 
Brentwood; 11 mi. south of Kettleman City; north of Davis, Yolo County; 
and Ventura. A record of "Compositae" near Los Angeles ( Fall, 1901 ) can 
probably be discounted. 

Subgenus POMPHOPOEA LeConte, new status 

Pomphopoea LeConte, 1862, Smithsonian Misc. Coll., vol. 3, art. 136, p. 
273; 1866, Smithsonian Misc. Coll., vol. 6, art. 167, p. 161. Horn, 1873, Proc. 
Amer. Philos. Soc, vol. 13, p. 115. Blatchley, 1910, The Coleoptera or beetles 
of Indiana, p. 1364. Cockerell and Harris, 1925, Proc. Biol. Soc. Washington, 
vol. 38, p. 32. Cros, 1929, Ann. Soc. Ent. France, vol. 98, pp. 213-14. Denier, 
1933, Rev. Chilena Hist. Nat., vol. 37, p. 241. Dillon, 1950, Amer. Midland 
Nat., vol. 48, p. 353. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, 
p. 71. 

Moderate-sized to large, slender beetles, most similar in general facies 
to members of the subgenus Lytta. Metallic green, blue, or aeneous with 
femora and tibiae (hind tibiae only in polita) orange. Head without 
a pale frontal spot. Pubescence colorless. At least elytra satiny in texture, 
not particularly shiny. Head with tempora well defined; surface micro- 
granulate, sparsely, finely punctate. Labrum deeply emarginate (Figs. 
155-58). Antennae moniliform clavate or (polita) subfiliform. Male 
antennae longer than in female; intermediate segments not incrassate or 



SYSTEMATICS AND PHYLOGENY 223 

otherwise modified. Pronotum subcampanuliform or subhexagonal. Wings 
medium brown. Under surface moderately densely pubescent; setae long 
except on midventral region of male abdomen, where they are much 
shortened. Fore femora at least slightly concave on anterior side so as to 
appear bowed, even in female. Male fore and middle legs modified or not. 
Male fore tibiae each with two spurs or only one (anterior spur absent 
in polita). Male middle tibiae each with posterior spur long, curving 
strongly posteriad. Tarsal pads well developed and pale on all segments, 
divided or parted on midline. Outer hind tibial spur expanded, flared; 
inner spur spiniform or sticklike, excavate behind. Male fifth abdominal 
sternum distinctly emarginate. Male genitalia with gonostyli elongate, 
slender, tapered, glabrous, blunt and turned dorsad at apex; aedeagus 
slender, not much curved; ventral hooks subequal in size and similar 
in shape, well separated; dorsal hook abruptly recurved. 

Type. Lytta sayi LeConte; fixed by subsequent designation by Mac- 
Swain (1956, p. 72, footnote). Dillon's (1952) designation of aenea as 
type is invalid because this species was not one of the first included in 
Pomphopoea. 

Remarks. Pomphopoea contains a total of four well-marked but super- 
ficially similar species, which fall naturally into three species groups. 
Adults of the species, in the main, are found in the spring on the blossoms 
of a variety of trees and shrubs, especially of members of the family 
Rosaceae. The range of the subgenus, which is centered in the Eastern 
Deciduous Forest, covers almost all of the eastern United States in addi- 
tion to a narrow, adjacent band in southeastern Canada (Fig. 9). The 
western limits of distribution are eastern Wyoming (sayi) and eastern 
Texas (aenea). As discussed in an earlier section of this paper, Pom- 
phopoea seems to have had a history of dispersal quite distinct from 
that of the other North American members of the genus Lytta, which it 
replaces east of the Great Plains. This geographical replacement cannot 
be explained entirely on a historical basis, however, for several of the 
primarily western species of Lytta have access to the Eastern Deciduous 
Forest at present or have undoubtedly had such access in relatively 
recent times. Rather, I would suggest that a critical deterrent to east- 
ward dispersal of members of the large western fauna of Lytta is an 
inability on their part to compete successfully with species of Pompho- 
poea in the mesic country east of the Great Plains. Similarly, westward 
movement of Pomphopoea is probably blocked by the presence of the 
western fauna of Lytta. 

Two of the species of the subgenus have ranges which are divided more 
or less completely into western and eastern segments. This segmentation 
of range presumably reflects post-Pleistocene changes of climate which 



224 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytttt 



CO 



o 

Q- 



< 
UJ 

z 

LU 

< 





or 




< 




_i 




3 




O 


.^ 


3 


>- 


(3 


< 


Z 


CO 


3 



Pol it a 
Group 



Aenea 
Group 



Sayi 
Group 



Fig. 60. Phylogeny of the subgenus Pomphopoea. 



have brought about partial or total extinction of the populations in the 
intermediate territory. Further study of the distribution and ecology 
of the members of the subgenus will probably lead to valuable conclu- 
sions regarding post-Pleistocene biogeography of the eastern United 
States. 

The phylogeny of the species of Pomphopoea, as suggested by the 
results of the present analysis of adult morphology, may be outlined as 
follows (see also Fig. 60). The subgenus originally arose from an an- 
cestral species which reached eastern North America from Asia by way 
of the Bering Strait and which was characterized, in all probability, 
by having the male fore tibiae not thickened or arcuate and with two 
spurs each, male hind tibiae not dilated (as in the female), gonostyli of 
the male genitalia parallel or nearly so, male sixth abdominal sternum 
simply emarginate, and female sixth sternum entire. All these characters 



SYSTEMATICS AND PHYLOGENY 225 

are now represented in the subgenus, although the above combination of 
them has not been retained by any single species. 

The species polita arose as an offshoot from the ancestral type in 
which in the male the fore tibiae developed a strong groovelike impres- 
sion, one of the fore tibial spurs was lost, the first fore tarsal segment was 
asymmetrically dilated, and the gonostyli became divergent. The sub- 
filiform nature of the antennae of this species, although it is unique within 
the subgenus, is perhaps a primitive rather than specialized character 
inasmuch as it approaches the condition in some of the Old World species 
of Lytta that I believe are closely related to Pomphopoea. Polita is ap- 
parently the least successful of the species of Pomphopoea, being both 
rare and confined in distribution. 

The other species of the subgenus apparently arose as derivatives of 
a separate evolutionary line, as indicated by the fact that in all of them 
the male hind tibiae are somewhat dilated and more strongly compressed 
than in the female and that the female sixth abdominal sternum is at 
least weakly emarginate. On this line aenea has retained an essentially 
primitive combination of characters, while in both sayi and unguicularis 
the male fore tibiae are thickened and arcuate, the male sixth abdominal 
sternum has become impressed, and the female sixth sternum is rela- 
tively deeply grooved. Unguicularis, with its strongly dilated and com- 
pressed male hind tibiae and strongly modified female sixth sternum, is 
clearly more highly evolved than sayi and is nearly if not fully as highly 
evolved as polita. It is of interest to note that while sayi and unguicularis 
replace each other geographically, the more distantly related aenea, 
which is the commonest and most widely distributed species of the 
subgenus, is able to occur in broad sympatry with both species. 

First instar larvae of sayi and aenea have been described by MacSwain 
(1956). These larvae show well-marked differences, but until the larva of 
at least one other species is known, their morphology cannot contribute 
to the study of phylogenetic relationships within the subgenus. 

Key to Groups 

1. Antennae subfiliform (Fig. 114); fore and middle tibiae black; male 
with a broad impression on anterior side of fore tibiae and a weaker 
one on middle tibiae; male fore tibiae each with only a single spur; 
first segment of male fore tarsi dilated on one side; male hind tibiae 
as in female, not dilated; female sixth abdominal sternum entire, not 

modified Polita Group ( p. 226 ) 

Antennae moniliform, clavate (Fig. 115); all tibiae orange, at least 
at middle; male fore tibiae and fore tarsi not modified as above; male 
fore tibiae each with two spurs; male hind tibiae at least weakly di- 
lated, more strongly compressed than in female; female sixth ab- 



226 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 

dominal sternum at least weakly emarginate medianly at apex 

( usually also with a groove ) 2 

2. Male fore and middle legs not modified as below, as in female; male 
sixth abdominal sternum (Fig. 272) neither impressed nor concavely 

folded; female sixth sternum not or only very feebly grooved 

* . . Aenea Group ( p. 228 ) 

Male fore femora noticeably more strongly impressed on anterior side 
than in female; male fore tibiae thickened, stouter than middle 
tibiae, bowed; male sixth abdominal sternum (Figs. 273-74) im- 
pressed or concavely folded, at least along emargination; female sixth 

sternum (Figs. 288-89) deeply grooved medianly at apex 

Sayi Group (p. 234) 

POLITA GROUP 

Head subtriangular; occiput prominent, well defined; surface of head 
and pronotum only moderately granulate, shiny. Antennae subfiliform. 
Pronotum subhexagonal, about as wide as long, the lateral angles better 
marked than in the Sayi and Aenea groups; disk bulged for basal two- 
thirds. Elytra very finely, obscurely rugose punctate, glabrous. Male fore 
tibiae each with only a single spur (anterior spur absent); anterior side 
of tibiae with a broad groovelike impression from before middle to apex, 
the impression forming a rather deep, impunctate concavity apically. 
Male middle tibiae with a weak groovelike impression subapically on 
lower half of anterior surface. Outer hind tibial spur twice as wide as 
inner spur, with excavation nearly longitudinal. Male fore tarsi with first 
segment asymmetrical, dilated on anterior side. Male sixth abdominal 
sternum neither impressed nor concavely folded. Female sixth sternum 
entire, not grooved. Male genitalia with gonostyli divergent apically. 
Female genitalia with anteromesal corner of each gonocoxite not pro- 
tuberant or obscuring notch of mesal margin. 

Remarks. This group contains only polita, from the southeastern United 
States. 

Lytta (Pomphopoea) polita Say 

Lytta polita Say, 1824, Jour. Acad. Nat. Sci. Philadelphia, vol. 3, p. 302. 
LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 336. 

Lytta femoralis LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 336. 

Pomphopoea polita, LeConte, 1866, Smithsonian Misc. Coll., vol. 6, no. 
167, p. 161. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 116. Sherman, 
1913, Ent. News, vol. 24, p. 247. 

Cantharis pedestris Gemminger, 1870, [Harold] Coleopt. Hefte, vol. 6, 
p. 124. New name for Lytta femoralis LeConte, not Klug, 1825, Nova Acta 



SYSTEMATICS AND PHYLOGENY 227 

Phys.-Med. Acad. Caesareae Leopoldino-Carolinae, vol. 12, pt. 2, p. 435. 

Pomphopoeca [sic] polita Hebard, 1903, Ent. News, vol. 14, p. 260. 

Potnphopoea femoralis, Blatchley, 1920, Canad. Ent., vol. 52, p. 72. 

Pomphopoea polita a. femoralis, Loding, 1945, Geol. Surv. Alabama, Mon. 
11, p. 55. 

Head, pronotum, and under surface aeneous, shiny. Elytra a duller 
brassy color. Femora orange with black apex. Hind tibiae orange with 
black base and apex. Fore and middle tibiae, and the tarsi, black. Black 
areas with a green luster. Length: 13 mm. -22 mm. 

Head subtriangular; surface sparsely punctate throughout, with very 
short, inconspicuous setae. Labral emargination extending to middle 
(Fig. 155). Pronotum usually with a pair of foveae on disk behind middle; 
surface of disk smooth, shiny, subimpunctate, glabrous. Tarsi with mid- 
line of ventral surface glabrous, impunctate, broadly dividing all tarsal 
pads. 

Male. Antennae as in Figure 114, only slightly longer than in female. 
Male fore tibiae with anterior, impressed surface very sparsely clothed 
with fine setae except for glabrous apical concavity; anteroventral 
margin of male fore tibiae fringed with setae, those at ventral corner 
of apex very dense, longer than rest, curving out over concavity. Fore 
tibial spur curving posteriad. Emargination of sixth abdominal sternum 
(Fig. 271) straight-sided; lateral lobes subacutely triangular. Genitalia 
as in Figure 348. 

Female. Antennae as in Figure 152. 

Type locality. Of polita, Georgia; of femoralis, Louisiana. 
Geographic distribution. Coastal Plain and Piedmont of the south- 
eastern United States from North Carolina to Louisiana. (See Fig. 61.) 
Seasonal distribution. December 19 — June 2. 

Records. UNITED STATES: Alabama: Auburn, June (Loding, 1945); 
Mobile County, February, March (Loding, 1945). Florida: Archbold 
Biological Station, Lake Placid, January, 3; Billy Island, Okefenokee 
Swamp, December, 5; Deland, March, 3; Dunedin, January (Blatchley, 
1920); Eau Gallie, March, 2; Hilliard, April, 2; Lake Placid, January, 1; 
Myakka River State Park, March, 2; Oneco, March, April, 28; Port Sewall, 
January, March, 3; Punta Gorda, February, March, 2; Tarpon Springs, 
March, 1. Georgia: Augusta, March, 1; Doughtry County, April, 1; Stone 
Mountain, DeKalb County, April, 1; Thomasville (Hebard, 1903); Way- 
cross, March, 8. Louisiana: (LeConte, 1853). Mississippi: Lucedale, 
February, 1. North Carolina: New River, April, May, 2; Raleigh, April, 
May, 2; Southern Pines, April, 1. South Carolina: (Horn, 1873). 

Remarks. The granulation of the surface of the head is usually more 



228 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 




Fig. 61. Distribution of Lytta polita. 



strongly marked in females than in males. The species is easily recognized 
by its aeneous color and subfiliform antennae. 

The type of polita has been lost. As neotype I have designated a 
male labeled Waycross, Georgia, March 22, 1952, J. R. McGillis, in the 
Canadian National Collection, at Ottawa. The type of femoralis is in 
Museum of Comparative Zoology. 

Bionomics. This species has been collected at light several times. No 
food plants are recorded. 



AENEA GROUP 

Head subtriangular; occiput not prominent. Antennae moniliform, 
clavate. Surface of head and pronotum strongly granulate, satiny. Pro- 
notum campanuliform, as in Sayi Group. Elytra rugose, sparsely pubes- 
cent to essentially glabrous. Male fore and middle legs not modified, as 
in female. Male hind tibiae moderately dilated. Male sixth abdominal 
sternum neither impressed nor concavely folded. Female sixth sternum 
weakly emarginate medianly at apex, not or only very feebly grooved. 



SYSTEMATICS AND PHYLOGENY 229 

Male genitalia with gonocoxites parallel. Female genitalia with antero- 
mesal corner of each gonocoxite not protuberant or obscuring notch of 
mesal margin. 

Remarks. The only species included in this group is aenea, which 
ranges through most of the eastern United States, as far west as eastern 
Texas. 

Lijtta (Pomphopoea) aenea Say 

Lytta aenea Say, 1824, Jour. Acad. Nat. Sci. Philadelphia, vol. 3, p. 301. 
LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 337. 

Cantharis nigricornis LeConte, 1847, Jour. Acad. Nat. Sci. Philadelphia, 
ser. 2, vol. 1, p. 90. 

Cantharis filiformis LeConte, 1847, Jour. Acad. Nat. Sci. Philadelphia, ser. 
2, vol. 1, p. 91. 

Lytta filiformis, LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, 
p. 337. 

Lytta tarsalis Bland, 1864, Proc. Ent. Soc. Philadelphia, vol. 3, p. 71. 
Borchmann, 1917, in Junk and Schenkling, Coleopterorum catalogus, pt. 69, 
p. 99. 

Pomphopoea texana LeConte, 1866, Smithsonian Misc. Coll., vol. 6, no. 
167, p. 161. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 117. Tucker, 
1906, Trans. Kansas Acad. Sci., vol. 20, pt. 1, p. 88. Dillon, 1952, Amer. 
Midland Nat., vol. 48, p. 354. New synonymy. 

Pomphopoea aenea, Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 117. 
Ulke, 1902, Proc. United States Nat. Mus., vol. 25, pp. 32, 54. Blatchley, 
1910, The Coleoptera or beetles of Indiana, p. 1364. Sherman, 1913, Ent. News, 
vol. 24, p. 247. Britton, 1920, Connecticut State Geol. and Nat. Hist. Surv. 
Bull. 31, p. 238. Mutchler and Weiss, 1924, New Jersey Dept. Agr. Circ. 76, 
p. 9. MacSwain, 1956, Univ. California Publ. Ent., vol. 12, p. 73, pis. 13, 15. 

Typically metallic aeneous green but varying to a nearly pure metallic 
green or blue. Elytra typically cupreous or cupreous green, rarely pur- 
plish; in the Texas race (see below) and in some specimens from the 
southeastern United States they are nearly pure green or (rarely) blue. 
Femora and tibiae orange, the femora often black at apex. Tarsi varying 
from orange to fuscous. Length: 9 mm. -16 mm. 

Head with surface finely, moderately densely punctate on front, 
sparsely punctate on vertex; punctures coarser than in the Sayi Group; 
pubescence moderately long, erect, conspicuous. Labral emargination 
deep, extending to middle (Fig. 156). Pronotum with surface and pubes- 
cence as on vertex, the center subimpunctate, nearly glabrous. Legs 
slender. Tarsal pads clearly, rather broadly divided except on first seg- 
ment of hind tarsi, where pad is merely parted; midline of tarsi impunc- 
tate. 

Male. Antennae (Fig. 116) nearly as in sayi, reaching base of prono- 
tum, obviously longer than in female; segments a little more elongate 



230 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

than in the Sayi Group. Hind tibiae weakly dilated. Fifth abdominal ster- 
num with lateral lobes obtusely rounded, not bulged. Sixth sternum as 
in Figure 272. Genitalia as in Figure 349. 

Female. Antennae as in Figure 153. Genitalia much as in polita. 

Type locality. Of aenea, Pennsylvania; of nigricornis, St. Louis, Mis- 
souri; of filiformis and texana, Texas; of tarsalis, Illinois. 

Geographic distribution. The range of this species appears to be di- 
vided into two major segments. One extends from New Hampshire south 
to Alabama and west to Missouri. The other includes central Oklahoma, 
eastern Texas, and northwestern Louisiana. (See Fig. 62.) 

Seasonal distribution. February 27 — July 6 (both extremes in Texas). 
Most records are in April and May. 

Records. UNITED STATES: Alabama: Cheaka State Park, April, 3; 
Montgomery, 1. Connecticut: Hamden, May, 1; New Haven, May, 9. 
District of Columbia: District label only, 1; Washington and 4.5 mi. 
southwest of Washington, April, 37. Georgia: Atlanta, April, 1; Barnes- 




Fig. 62. Distribution of Lytta aenea. 



SYSTEM ATICS AND PHYLOGENY 231 

ville, March, 2; Camp McClellan, April, 1; Fort Valley, April, 1; Kenne- 
saw, March, 1; Marshalville, 3; Meriwether County, April, 1; Spalding 
County, 1; Stone Mountain, DeKalb County, March, April, 7. Illinois: 
Kickapoo State Park, April, 1; Mahomet, April, 1; Paris, April, 1; Put- 
nam County, May, 1; Savanna, May, 1. Indiana: Brown County, April, 
1; Lafayette, April, 1; Putnam County, April, 1; Vigo County, 1. Ken- 
tucky: State label only, 1; West Point, April, 1. Louisiana: State label 
only, 1; Shreveport, March, 10. Maryland: State label only, 1; Alberton, 
April, 5; Camp Springs, May, 1; Clear Spring, April, 12; College Park, 
1; Hyattsville, May, 1; Jenny, April, 1; Linden, April, 1; Plummers Island, 
April, 7; Takoma Park, May, 2. Massachusetts: Charlemont, 1; Cheshire, 
1; Hopkinton, May, 1; Milton, May, 2; Natick, May, 2. Missouri: State 
label only, 3; Cape Girardeau, April, 1; Charleston, April, 2; Columbia, 
April, May, 3; Ranken, April, 1; St. Louis (LeConte, 1847). New Hamp- 
shire: State label only, 7. New Jersey: Anglesea (Mutchler and Weiss, 
1924); Fort Dix, May 1; Glassboro, April, 5; Greenwood Lake, May, 1; 
Merchantville (Mutchler and Weiss, 1924); Trenton, May, 1; Westville 
and Woodside (Mutchler and Weiss, 1924). New York: Brewster, April, 
1; Hugenot Woods, Westchester County, May, 1; Ithaca, May, June, 2; 
Montgomery, May, 2; Mount Maclntyre, June, 1; New Windsor, April, 
May, 6; New York, May, 1; Rock City, May, 1; West Point, April, 3; 
Wilmington, June, 1. North Carolina: Ellenboro, March, 7; Faison, 
April, 1; Raleigh, April, 3; Waccamaco Lake, April, 1; Whiteville, April, 
1; Wilmington, April, 1. Ohio: Chestnut Grove, Ashtabula County, June, 
1; Cincinnati, April, May, 4; Springfield, April, 1; Summit County, April, 
1. Oklahoma: Kingfisher, March, 3; Pawnee, March, 2; Payne County, 
April, 1. Pennsylvania: Allegheny County, April, May, 4; Cabot, April, 2; 
Danville, May, 1; Darby, April, 1; Germantown, May, 1; Hazleton, 1; 
Jeannette, 1; Philadelphia, April, May, 4; Reading, April, 2; Washington 
County, 1; Wilawana, April, June, 3; Windgap, May, 1; Wyoming, April, 
1. South Carolina: Clemson College, April, 1; Conway, March, 1. Ten- 
nessee. State label only, 1; Great Smoky Mountains, 1600 ft., April, 3; 
Knoxville, April, 1; Reelfoot Lake, 1; Tazewell, April, 2. Texas: State label 
only, 41; Austin, May, 1; Bethage, 1; Brazos County, April, 2; Bryan, 
March, 1; College Station, February, April, 2; Colorado County, March, 
1; Comanche County, March, 1; Dallas, March, 3; Dallas County, July, 1; 
Fedor, 5; Fort Worth, 2; Gatesville, April, 1; Iowa Park, March, 1; Mason, 
May, 1; Ranger, March, 6; San Antonio, March, 3; Thornberry (Tucker, 
1906); Tiger Mills, April, 2. Virginia: Falls Church, April, 10; Great Falls, 
April, 2; Ivy Depot, April, 3; Rosslyn, April, 1; Salem, 1; Vienna, April, 
3; West Falls Church (MacSwain, 1956). West Virginia: Levels, April, 4. 
Remarks. Geographic variation in this species is summarized in 
Tables VII and VIII. For purposes of analysis of variation the specimens 






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SYSTEMATICS AND PHYLOGENY 233 

Table VIII. Degree of Dilation op Male Hind Tibia of Lytta aenea 
Expressed as Average Ratio of Greatest Width of Tibia to Least Width at Base 





average 


STANDARD 


STANDARD 


NUMBER OF 




ratio 


ERROR 


DEVIATION 


SPECIMENS 


New England, New Jersey, 










eastern Pennsylvania 


1.73 


.03 


.13 


24 


Ohio, Indiana, Illinois, western 










Pennsylvania, Kentucky 


1.76 


.03 


.09 


8 


Missouri 


1.77 






3 


Maryland, District of Columbia 


1.86 


.02 


.12 


26 


Virginia, West Virginia 


1.86 


.04 


.14 


11 


North Carolina, South Carolina, 










eastern Tennessee 


2.01 


.06 


.16 


8 


Alabama, Georgia 


1.98 


.04 


.16 


15 


Texas, Oklahoma, Louisiana 


2.17 


.03 


.19 


32 



examined have been divided into eight geographic samples, as indicated 
in the tables. The main variational trends are discussed briefly below. 

As shown in Table VII, the femora are frequently tipped with black 
in specimens from northern populations, while they are not or only 
very rarely so marked in specimens from the southern part of the range. 
Color of the tarsi varies discordantly. For the most part the frequency of 
occurrence of black or fuscous coloring of the tarsi is comparable to 
that of black tipping of the femora, but major exceptions occur in the 
Maryland-District of Columbia population and even more strikingly in 
the Texas-Oklahoma-Louisiana population, where all specimens have 
dark tarsi in the complete absence of black femoral tipping. 

The distinctness of the latter population is also apparent with respect 
to color of the elytra. Although the data are not given in Table VII, the 
color of the head, pronotum, and under surface varies also. In general 
these parts are an aeneous or olive green in specimens with at least some 
cupreous tone in the elytra and a more nearly pure green or blue in 
specimens with green or blue elytra. It should be noted that color varia- 
tion in aenea is continuous in nature, and that the three classes of elytral 
color recognized in Table VII are arbitrary. 

Density of pubescence varies greatly, with a strong tendency for 
denser development in northern populations. A subtle but distinct north- 
south trend for increased dilation of the male hind tibiae is present, as 
indicated by the data given in Table VIII. 

The distributional data (Fig. 62) suggest that the Texas-Oklahoma- 
Louisiana population of aenea may be geographically isolated from more 
northern and eastern populations. At the same time, the somewhat 



234 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

intergradational nature of the Alabama-Georgia population would indi- 
cate that if isolation is indeed a fact, it is of relatively recent origin. To 
the west, there is apparently no gene exchange between the southwestern 
population and the Missouri population. 

On the basis of the information now available, I will speculate that 
during the last glacial epoch the range of aenea was broadly continuous 
through the southeastern United States and that as the species moved 
northward with the subsequent amelioration of climatic conditions con- 
tact with the southwestern population was lost or severely limited. The 
absence of intergradation between the southwestern population and the 
Missouri population may be explained on the basis that the latter pushed 
into Missouri from Illinois and has never reached far enough west to 
contact the population in Oklahoma. 

Separation of the southwestern population of aenea as a separate 
species (texana) does not seem justifiable on a morphological basis, 
although there is ample reason to recognize this population as a separate 
race. A vernacular name, such as "Texas race," would seem appropriate 
for it. 

The first instar larva of aenea has been described by MacSwain ( 1956). 

As neotype of aenea I have designated a male in the United States 
National Museum from Allegheny County, Pennsylvania (Wickham col- 
lection ) . The types of the synonyms nigricornis, filiformis, and texana are 
in the Museum of Comparative Zoology. The type of tarsalis is in the col- 
lection of the Academy of Natural Sciences of Philadelphia. 

Bionomics. This species appears to be similar to sayi in its feeding 
habits. Genera of Rosaceae on whose flowers it is known to feed include 
Amelanchier, Crataegus, Prunus (peach, plum), Pirophorum (pear) 
(recorded eating fruit also by Mutchler and Weiss, 1924), and Mains 
( apple ) . In addition, there are single records each of feeding on flowers 
of willow (Salicaceae) (Ulke, 1902) and Carya (Juglandaceae) and 
leaves of oak (Fagaceae) (Sherman, 1913). 

SAYI GROUP 

Head triangular; occiput not very prominent. Surface of head and 
pronotum strongly granulate, satiny. Antennae moniliform clavate. Pro- 
notum campanuliform, about as wide as long, impressed at base medianly; 
disk even, rather flat. Elytra distinctly rugose punctate, essentially gla- 
brous. Male fore femora very noticeably more strongly impressed on 
anterior side than in female; anterior surface smoother and more nearly 
glabrous (setae minute, erect, very sparse). Male fore tibiae thickened, 
stouter than middle tibiae, decidedly bowed; anterior surface like that 
of fore femora. Male hind tibiae at least weakly dilated. Outer hind tibial 
spur less obliquely excavate than in the Polita Group. Male sixth abdomi- 



SYSTEMATICS AND PHYLOGENY 235 

nal sternum impressed, at least along emargination. Female sixth sternum 
notched and deeply grooved medianly at apex. Male genitalia with gono- 
styli parallel. Female genitalia with anteromesal corner of each gonocoxite 
forming a dark, protuberant tooth which obscures notch of mesal margin 
in ventral view; notch weak. 

Remarks. This group includes the species sayi and unguicularis. Its 
range encompasses a large part of the eastern United States and a narrow, 
adjacent area of Canada. 

Key to Species 

1. Tibiae narrowly black at base and apex; labral emargination not ex- 
tending to middle ( Fig. 157 ) ; male fore femora not contorted; female 
sixth abdominal sternum with groove not strictly vertical in position 

(Fig. 288) .' sayi (p. 235) 

Tibiae entirely orange; labral emargination deeper, extending at least 
to middle (Fig. 158); male fore femora contorted; female sixth ab- 
dominal sternum with groove strictly vertical in position 

unguicularis ( p. 238 ) 

Lytta (Pomphopoea) sayi LeConte 

Lytta aenea, Say, 1824, in Keating, Narrative of an expedition to the source 
of St. Peters River . . . , vol. 2, p. 288. Misidentification. 

Lytta sayi LeConte, 1853, Proc. Acad. Nat. Sci. Philadelphia, vol. 6, p. 336. 

Cantharis pyrivora Fitch, 1859, Third report on the noxious and other in- 
sects of the State of New York, p. 36. 

Pomphopoea sayi, LeConte, 1866, Smithsonian Misc. Coll., vol. 6, no. 167, 
pp. 160-61. Horn, 1873, Proc. Amer. Philos. Soc, vol. 13, p. 117. Wickham, 
1896, Canad. Ent, vol. 28, p. 34. Blatchley, 1910, The Coleoptera or beetles 
of Indiana, p. 1364, fig. 590. Gibson, 1912, 42nd Ann. Rept. Ent. Soc. Ontario, 
p. 4, fig. 7. Brirton, 1920, Connecticut State Geol. and Nat. Hist. Surv. Bull. 
31, p. 238. Mutchler and Weiss, 1924, New Jersey Dept. Agr. Circ. 76, p. 9. 
Cockerell and Harris, 1925, Proc. Biol. Soc. Washington, vol. 38, p. 31, 
fig. 10. Houghton, 1947, Bull. Brooklyn Ent. Soc, vol. 42, pp. 103-05. Mac- 
Swain, 1956, Univ. California Publ. Ent., vol. 12, p. 72, pi. 14. 

Metallic green or blue. Femora orange with black apex. Tibiae orange 
with black base and apex. Tarsi black. Black areas with a green or blue 
luster. Length: 13 mm. -22 mm. 

Head with surface finely, sparsely punctate; pubescence moderately 
long, conspicuous. Labral emargination not extending to middle (Fig. 
157). Pronotum with surface and pubescence as on vertex, the center 
subimpunctate, nearly glabrous. Tarsal pads narrowly divided, those of 
basal segments often only parted; midline of tarsi punctate. 

Male. Antennae as in Figure 117, reaching base of pronotum, a little 
longer than in female. Fore femora not contorted. Hind tibiae weakly 



236 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

dilated. Fifth abdominal sternum with lateral lobes obtusely rounded, 
not bulged. Sixth sternum (Fig. 273) impressed along emargination, not 
folded; lateral lobes prominent, evenly rounded, obtuse. Genitalia as in 
Figure 350. 

Female. Antennae as in Figure 154. Sixth abdominal sternum more 
deeply notched than in unguicidaris and with groove less vertical and 
extending farther basad (Fig. 288). 

Type locality. Of sayi Illinois; of pyrivora, Canajoharie, New York. 

Geographic distribution. Southern Quebec and Pennsylvania to north- 
eastern Wyoming and Iowa. (See Fig. 63.) Available distributional data 
suggest that there are three isolated populations. 

Seasonal distribution. May 18 — July 27. The vast majority of the records 
are in May and June. Three specimens examined are labeled as collected 
in September. 

Records. CANADA: Ontario: Brittania, May, 4; Hastings, May, 1; 
Hastings County, May, 1; Kearney, July, 2; Kinburn, June, 7; Marmora, 
May, 13; Merivale, June, 1; Muskoka (Gibson, 1912); Osgoode [Sta- 
tion], May, June, 3; Renfrew, 1; Smoky Falls, Mattagami River, June, 1; 
Perkins, May, 1; Sudbury, 2; Toronto (Gibson, 1912); Westboro, June, 
9; Wright, May, 2. Quebec: Beech Grove, June, 1; Covey Hill, June, 1; 
Duparquet, May, 14; Harrington Lake, Gatineau Park, June, 1; Hull 
( Gibson, 1912 ) ; Kazubazua, June, 2; Showbridge ( Gibson, 1912 ) ; Wake- 
field, June, 1. 

UNITED STATES: Connecticut: Cornwall, June, 18. Illinois: Freeport, 
May, 3; Morrison, June, 12; Oregon, May, June, 3; Savanna, 1; Urbana, 
June, 1. Iowa: Benton County, June, July, 2; Delaware County, June, 1; 
Des Moines, June, 2. Massachusetts: Hopkinton, June, 1; Montgomery, 
June, 1; Mount Everett, June, 2; Mount Wachisett, 1; Williamstown, June, 
1. Minnesota: Itasca State Park, June, 18. New Jersey: Greenwood Lake, 
June, 1. New York: Albany, May, 1; Almond, June, 6; Barre, June, 1; 
Cambridge, June, 1; Canajoharie, June, 1; Canandaigua, 1; Canton, June, 
1; Castleton, June, 7; Clinton, June, 2; Connecticut Hill, Tompkins 
County, June, 2; Cooks Falls, 1; Corinth, June, 1; Elba, 1; Elbridge, May, 
2; Ellenberg, June, 11; Elsmere, May, 2; Fonda, June, 2; Genesee County, 
June, 4; Geneva (MacSwain, 1956); Gloversville, May, June, 14; Hamil- 
ton, June, 1; Hartford, May, 15; Hudson, May, 1; Ithaca, May, June, 18; 
Lockport, June, 1; McLean, 11; Meadowdale, May, 1; Newport, June, 
1; Northampton, June, 1; North Elba, July, 3; North Fairhaven, June, 1; 
North Java, July, 3; Patterson, June, 4; Peru, June, 1; Philmont, June, 1; 
Portage, 1; Port Jervis, June, 3; Poughkeepsie, June, 1; Putnam, June, 2; 
Ravena, June, 1; Rochester, June, 1; Schenectady County, June, 1; Schuy- 
lerville, June, 1; Slingerland, May, 5; Sonyea, July, 4; Tuscarora, 1; 



SYSTEMATICS AND PHYLOGENY 



237 



Varna, May, 1; Voorheesville, June, 4; Waterville, June, 1; Waver ly, June, 
3; Wellsburg, July, 2; Wilmurt, 2. Pennsylvania: Factory ville, June, 1; 
Hummelstown, 3 ( 2 labeled September ) ; Mount Pocono, July, 7; Muney, 
June, 1; Sylvania, 1; Wilawana, June, 8; Wysox, June, 2. Vermont: Mid- 
dleburg, May, 3. Wisconsin: Cedar Lake, June, 1; Dane County, Sep- 
tember, 1. Wyoming: Crook County, June, 6. 

Remarks. Material from Ontario and Quebec is blue or (less fre- 
quently) greenish blue in color except for a series from Beech Grove, 
Quebec, which is a rather pure green. In the United States the blue color 
is characteristic of samples of sayi from New York and eastward, although 
within this area color varies in series through blue-green to an occasional 
pure green. The Canadian and eastern United States material presum- 
ably represents a single population. In Illinois, Wisconsin, Minnesota, and 
Iowa there is a second population characterized by a purer, brighter 
green color, with a tendency for the development of a brassy quality. The 
Wyoming series, representing a third population, has a dark bluish green 
color quite distinct from that of either the eastern or the central popula- 
tion. In addition, it is unusual in that the orange of the legs ( especially the 




Fig. 63. Distribution of Lytta sayi (circles) and Lytta unguicularis (squares). 



238 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

hind ones) is darkened to a rather dull piceous. I have found no signifi- 
cant structural variation in the species. 

The first instar larva of sayi has been described by MacSwain (1956). 
The type of sayi is in the Museum of Comparative Zoology. The type of 
pyrivora has been lost. 

Bionomics. Adults have been recorded in the literature from the follow- 
ing plants : Prunus ( peach, cherry, plum ) , Pirophorum ( pear ) , and Rosa 
( Roseaceae ) ; Kolkwitzia, elder, and Viburnum lentago ( Caprifoliaceae ) ; 
Robinia pseudo-acacia and beans (Leguminosae); butternut (Juglanda- 
ceae); and wheat (Gramineae) (Fitch, 1859; Gibson, 1912; Houghton, 
1947; and Mutchler and Weiss, 1924). Some of the specimens examined 
from Ontario, Canada, are labeled as collected on Salix discolor ( Salica- 
ceae) and Cornus (Cornaceae). The beetles apparently feed largely on 
flowers; Fitch reported that they also attack young fruit of Rosaceae. 
Houghton stated that each year over a period of seven years a swarm of 
hundreds of the beetles arrived at a garden in Springville, New York, 
just as two large Robinia trees were flowering. After stripping these trees 
of flowers, the swarm began feeding on the flowers of nearby Kolkwitzia 
and Rosa shrubs. Two or three days after arrival it moved on. 

Lytta (Pomphopoea) unguicularis (LeConte) 

Pomphopoea unguicularis LeConte, 1866, Smithsonian Misc. Coll., vol. 6, 
no. 167, p. 160. Horn, 1873, Proc. Amer. Philos. Soc., vol. 13, p. 116. Blatchley, 
1910, The Coleoptera or beetles of Indiana, p. 1364. Sherman, 1913, Ent. 
News, vol. 24, p. 247. Britton, 1920, Connecticut State Geol. and Nat. Hist. 
Surv. Bull. 31, p. 238. 

Metallic green. Femora, tibiae, and tarsi orange, the femora narrowly 
black at apex. Length: 17 mm.-25 mm. 

Head and pronotum essentially as in sayi except that labral emargina- 
tion is deeper, extending at least to middle ( Fig. 158 ) . Tarsi with midline 
of ventral surface glabrous, impunctate, broadly dividing all tarsal pads. 

Male. Antennae as in sayi. Fore femora deeply concave on anterior 
side, contorted (twisted 45 degrees anteriad). Hind tibiae strongly di- 
lated and compressed, at apex as wide or nearly as wide as middle of 
femora. Fifth abdominal sternum with lateral lobes conspicuously bulged. 
Sixth sternum as in Figure 274, very deeply impressed ( concavely folded 
in most dried specimens). Genitalia as in sayi. 

Female. Antennae and genitalia essentially as in sayi. Sixth abdominal 
sternum with notch shallower than in sayi; groove shorter, strictly ver- 
tical in position (Fig. 289). 

Type locality. Illinois. 

Geographic distribution. Eastern Alabama northwest to Illinois and 
northeast to the Smoky Mountains. (See Fig. 63.) 



SYSTEMATICS AND PHYLOGENY 239 

Except for a record from Connecticut, the range of iinguicularis is 
allopatric with that of sayi. The species was reported from Connecticut 
by Britton (1920), who mentioned no specific locality. Although I have 
seen no specimens on which inclusion of the species in the Connecticut 
list could have been based, I have examined a series of 23 specimens 
labeled Cornwall, Connecticut, June 16, 1922 (2 labeled June 20), Cham- 
berlain collection (New York University, Cornell University, and United 
States National Museum ) . However, under the circumstances ( including 
the lack of collector's name on the labels), I strongly suspect the validity 
of the locality data, and I think the occurrence of iinguicularis in Con- 
necticut needs verification. 

Seasonal distribution. May 2 — July 4. 

Records. UNITED STATES: Alabama: Auburn, May, 1. Illinois: (Le- 
Conte, 1866). North Carolina: Black Mountain, May, 1; Blowing Rock 
(Sherman, 1913); Montreat, July, 3; Mount Graybeard, May, June, 3; 
Mount Mitchell, 6711 ft., June, 1. Tennessee: Great Smoky Mountains, 
June, 58; Matin, June, 1. 

Remarks. The type is in the Museum of Comparative Zoology. 

Rionomics. There are records of adults feeding on both Rosaceae and 
Ericaceae. Sherman (1913) reported that in June 1901 at Blowing Rock, 
North Carolina, adults swarmed in "untold thousands" on peach, rose, 
and mountain laurel (Kalmia). According to his account, they ate the 
blossoms of the mountain laurel and the leaves of the peach, "apparently 
preferring those [leaves] that were affected with leaf curl disease"; the 
feeding habits on rose were not mentioned. The specimens examined from 
the Great Smoky Mountains, Tennessee, were collected on azalea (Rho- 
dodendron). 



Literature Cited in the Text 

Audouin, V. 

1826. Prodrome d'une histoire naturelle des Cantharides. Ann. Sci. Nat., 
pp. 31-62, pi. 9. 
Beauregard, H. 

1885. Sur le mode de developpement naturel de la cantharide. Comptes 
Rendus Seances Acad. Sci. Paris, 1885, pp. 1472-75. 

1889. [Notes on Meloidae.] Bull. Soc. Ent. France, 1889, pp. ccxii-ccxiii. 

1890. Les insectes vesicants. Paris. 
Blackwelder, R. E. 

1939. Fourth supplement, 1933 to 1938 (inclusive), to the Leng catalogue of 
Coleoptera of America, north of Mexico. Mount Vernon, New York. 

1945. Checklist of the coleopterous insects of Mexico, Central America, the 
West Indies, and South America. Part 3. U.S. Nat. Mus. Bull. 185. 
Blatchley, W. S. 

1920. Notes on the winter Coleoptera of western and southern Florida, with 
descriptions of new species. Canad. Ent., vol. 52, pp. 68-72. 

BORCHMANN, F. 

1917. Pars 69: Meloidae, Cephaloidae. In Junk, W., and S. Schenkling, 
Coleopterorum catalogus. Berlin. 
Britton, W. E. 

1920. Checklist of the insects of Connecticut. State Geol. and Nat. Hist. 
Surv. Bull. 31. 

Carpenter, G. D. H. 

1921. Experiments on the relative edibility of insects, with special reference 
to their colouration. Trans. Ent. Soc. London, 1921, pp. 1-105. 

Carruth, L. A. 

1931. The Meloidae of South Dakota (Coleoptera). Ent. News, vol. 42, pp. 
50-55. 
Champion, G. C. 

1891-93. Family Meloidae. In Godman, F. E., and O. Salvia, Biologia Cen- 
trali-Americana, Coleoptera, vol. 4, pt. 2, pp. 364-450, 462-64, pis. 17-21. 
London. 
Chevrolat, A. 

1834-35. Coleopteres du Mexique. Strasbourg. 
Chittenden, F. H. 

1899. Insects injurious to beans and peas. In Yearbook of the U.S. Depart- 
ment of Agriculture for 1898, pp. 250-51. Washington. 
1903. A brief account of the principal insect enemies of the sugar beet. U.S. 

Dept. Agr. Div. Ent. Bull. 43. 
1903. Some insects recently injurious to truck crops. U.S. Dept. Agr. Div. 
Ent. Bull. 40 (n.s.), pp. 114-20. 

COCKERELL, T. D. A. 

1898. Life zones in New Mexico. II. The zonal distribution of Coleoptera. 

New Mexico Agr. Exp. Sta. Bull. 28, pp. 137-79. 
1901. Flower and insect records from New Mexico. Ent. News, vol. 12, pp. 

38-43. 

240 



LITERATURE CITED IN THE TEXT 241 

Cushny, A. R. 

1901. A textbook of pharmacology and therapeutics. Philadelphia and New 
York. 
Denier, P. C. L. 

1940. Enumerationi Coleopterorum Americanorum familiae Meloidarum. 

Corrigenda et addenda. Rev. Soc. Ent. Argentina, vol. 10, pp. 418-25. 
1940. Desicrption [sic] de Lytta neivai, n. sp., du Bresil et notes sur quelques 
Lytta F. de l'Amerique du Sud (Col. Meloidae). Rev. Ent., vol. 11, pp. 
799-802. 
Dillon, L. S. 

1952. The Meloidae (Coleoptera) of Texas. Amer. Midland Nat., vol. 48, 
pp. 330-420. 
Duges, E. 

1869-81. Description de algunos Meloideos indigenas. La Naturaleza, vol. 1, 
pp. 100-113, 125-28, 157-68, pis. 1-2 (1869); vol. 4, pp. 57-67, pi. 2 
( 1877) ; vol. 5, pp. 140-48, pi. 4 ( 1881 ) . 
1889. Sinopsis de la Meloideos de la Republica Mexicana. An. Mus. Micho- 
acano, vol. 2, pp. 34-40, 49-114 [49-128]. 
Edwards, J. G. 

1951. [Note on Poreospasta polita Horn mating with Lytta auriculata Horn.] 
Pan-Pacific Ent., vol. 27, p. 44. 
Enns, W. R. 

1956. A revision of the genera Nemogiwtha, Zonitis, and Pseudozoniti.s 
(Coleoptera, Meloidae) in America north of Mexico, with a proposed new 
genus. Univ. Kansas Sci. Bull., vol. 37, pp. 685-909. 

ESCHERICH, K. 

1895. Beitrage zur Naturgeschichte der Meloidengattung Lytta Fab. Ver- 
handl. Kaiserlich-Koniglichen Zool.-Bot. Ges., vol. 44 (1894), pp. 251-98, 
pis. 8-11. 
Essig, E. O. 

1926. Meloidae. In Insects of western North America, pp. 389-91. New York. 
Fabre, J. H. 

1886. Souvenirs entomologiques. Troisieme serie. Paris. 
Fall, H. C. 

1901. Notes on Dichelonycha and Cantharis, with descriptions of new species 

in other genera. Trans. Amer. Ent. Soc, vol. 27, pp. 277-310. 
1901. List of the Coleoptera of southern California. Occas. Papers California 
Acad. Sci., vol. 8, pp. 1-282. 
Fall, H. C, and T. D. A. Cockerell. 

1907. The Coleoptera of New Mexico. Trans. Amer. Ent. Soc, vol. 33, pp. 
145-272. 
Fitch, A. 

1859. Third report on the noxious and other insects of the State of New York. 
Albany, New York. 
Fox, W. B. 

1943. Some insects infesting the "selenium indicator" vetches in Saskatche- 
wan. Canad. Ent., vol. 75, pp. 206-7. 
Fumouze, A. 

1867. De la cantharide officinale. Paris. 
Gibson, A. 

1912. Blister beetles. 42nd Ann. Rept. Ent. Soc Ontario, 1911, pp. 1-6 
[reprint pagination]. 



242 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

GOEDART, J. 

1700. Metamorphoses naturelles ou histoire des insectes. Paris. 
Goodman, L., and A. Gilman. 

1941. The pharmacological basis of therapeutics. New York. 
Gornitz, K. 

1937. Cantharidin als Gift und Anlockungsmittel fvir Insekten. Arb. Physiol. 
Angew. Ent. Berlin, vol. 4, pp. 116-57. 
Haag-Rutenberg 

1880. Beitrage zur Kenntniss der Canthariden. Deutsche Ent. Zeitschr., vol. 
24, pp. 17-90. 
Hebard, M. 

1903. Notes on the collecting around Thomasville, Georgia. Ent. News, vol. 
14, pp. 260-61. 
Hecht, M. K., and D. Marien. 

1956. The coral snake mimic problem: a reinterpretation. Jour. Morph., vol. 
98, pp. 335-56. 
Herrera, A., and G. Mendoza. 

1866. Apuntes para la monografia de los insectos vexicantes indigenas. 
Gaceta Med. Mexico, vol. 2, pp. 264-66, 1 pi. 
Hicks, E. A. 

1959. Checklist and bibliography on the occurrence of insects in birds' nests. 
Ames, Iowa. 
Horn, G. H. 

1868. New species of Coleoptera from the Pacific district of the United 

States. Trans. Amer. Ent. Soc, vol. 2, pp. 129-40. 
1870. Contributions to the coleopterology of the United States. Trans. Amer. 

Ent. Soc, vol. 3, pp. 69-97. 
1873. Revision of the species of several genera of Meloidae of the United 
States. Proc. Amer. Philos. Soc, vol. 13, pp. 88-117. 

1873. Coleoptera. In Rept. U.S. Geol. Surv. 1872, 1873, p. 717. 

1874. Descriptions of new species of United States Coleoptera. Trans. Amer. 
Ent. Soc, vol. 5, pp. 20-43. 

1885. Studies among the Meloidae. Trans. Amer. Ent. Soc, vol. 12, pp. 107- 

16, pi. 5. 
1894. The Coleoptera of Baja California. Proc. California Acad. Sci., vol. 4, 

pp. 302-449, pis. 7-8. 
1896. Coleoptera of Baja California. Supplement II. Proc. California Acad. 

Sci., vol. 6, pp. 367-81. 

HORSFALL, W. R. 

1941. Biology of the black blister beetle (Coleoptera: Meloidae). Ann. Ent. 
Soc. America, vol. 34, pp. 114-26. 

1942. Biology of the squash blister beetle. Jour. Kansas Ent. Soc, vol. 15, 
pp. 93-99. 

1943. Biology and control of common blister beetles in Arkansas. Univ. 
Arkansas Agr. Exp. Sta. Bull. 436, pp. 1-55. 

Houghton, F. 

1947. Say's blister beetles. Bull. Brooklyn Ent. Soc, vol. 42, pp. 103-5. 
Houlbert, C, and L. Betis. 

1913. Faune entomologique Armoricaine, Coleopteres, 65'' famille, Meloides 
(Vesicants). Paris. 



LITERATURE CITED IN THE TEXT 243 

Jaeger, E. C. 

1938. The California deserts. 2d ed. Stanford, California. 
Kaszab, Z. 

1959. Phylogenetische Beziehungen des Fliigelgeaders der Meloiden (Cole- 
optera), nebst Beschreibung neuer Gattungen und Arten. Acta Zool. Acad. 
Sci. Hungaricae, vol. 5, pp. 67-114. 
Knowlton, G. F. 

1930-31. Notes on Utah Coleoptera. Florida Ent., vol. 14, pp. 36-37, 53-56, 
75-77; vol. 15, p. 10. 

1939. Utah Coleoptera. Utah Agr. Exp. Sta. Mimeo. Ser. 200 (Tech.), pt. 3. 
Knowlton, G. F., and G. P. Taylor. 

1952. Beetles — Coleoptera. Records and notes, largely from Utah. Utah Agr. 

Exp. Sta. Mimeo. Ser. 389. 
Lampa, S. 

1903. Berattelse till Kongl. Landtbruksstyrelsen Angaende verksamheten 

vid Statens entomologiska Anstalt under Ar 1902. Ent. Tidskrift, pp. 1-60. 
LeConte, J. L. 

1847. Fragmenta entomologica. Jour. Acad. Nat. Sci. Philadelphia, ser. 2, 

vol. 1, pp. 71-93. 
1851. Descriptions of new species of Coleoptera, from California. Ann. Lye. 

Nat. Hist. New York, vol. 5, pp. 125-216. 
1853. Descriptions of some new Coleoptera from Texas, chiefly collected by 

the Mexican Boundary Commission. Proc. Acad. Nat. Sci. Philadelphia, 

vol. 6, pp. 429-48. 

1857. Report upon the insects collected on the survey. In Reports of explora- 
tions and surveys to ascertain the most practicable and economical route 
for a railroad from the Mississippi River to the Pacific Ocean made under 
the direction of the Secretary of War, in 1853-5, vol. 12, book 2, part 3, 
pis. 1-2. Washington. 

1858. Description of new species of Coleoptera, chiefly collected by the 
United States and Mexican Boundary Commission, under Major W. H. 
Emory, U.S.A. Proc. Acad. Nat. Sci. Philadelphia, 1858, pp. 59-89. 

1861. New species of Coleoptera inhabiting the Pacific district of the United 

States. Proc. Acad. Nat. Sci. Philadelphia, 1861, pp. 338-59. 
1863-66. New species of North American Coleoptera. Part I. Smithsonian 

Misc. Coll., vol. 6, no. 167, pp. 1-86 (1863), 87-177 (1866). 
1866. List of Coleoptera collected near Fort Whipple, Arizona, by Dr. Elliott 

Coues, U.S.A., in 1864-65. Proc. Acad. Nat. Sci. Philadelphia, 1866, pp. 

348-49. 
1878. The Coleoptera of the alpine regions of the Rocky Mountains. Bull. 

U.S. Geol. Sun^., vol. 4, pp. 447-80. 
Lichtenstein, J. 

1875. [Miscellaneous notes.] Bull. Soc. Ent. France, 1875, pp. CLVI-CLVIII, 
CLXIII-CLXIV, CCI-CCII. 

1877. Quelques feuillets de mon journal. Mittheilungen Schweizerischen 
Ent. Ges., vol. 5, pp. 297-303. 

1878. [Miscellaneous observations.] Bull. Soc. Ent. France, 1878, pp. 
LXXXIX-XC. 

1879. [Miscellaneous notes.] Bull. Soc. Ent. France, 1879, pp. XXIV-XXVI, 
LXV, LXXII, LXXVI. 



244 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

Linsley, E. G., and J. W. MacSwain. 

1942. The parasites, predators, and inquiline associates of Anthophora lins- 
leyi. Amer. Midland Nat., vol. 27, pp. 402-17. 

1952. Notes on some effects of parasitism upon a small population of Diada- 
sia bituberculata (Cresson) (Hymenoptera: Anthophoridae). Pan-Pacific 
Ent, vol. 28, pp. 131-35. 
1958. The nesting habits, flower relationships, and parasites of some North 
American species of Diadasia (Hymenoptera: Anthophoridae). Wasmann 
Jour. Biol., vol. 15, pp. 199-235. 
Linsley, E. G., J. W. MacSwain, and R. F. Smith. 

1956. Biological observations on Ptilothrix sumichrasti (Cresson) and some 
related groups of emphorine bees (Hymenoptera, Anthophoridae). Bull. 
Southern California Acad. Sci., vol. 55, pp. 83-101. 
Loding, H. P. 

1945. Catalogue of the beetles of Alabama. Geol. Surv. Alabama, Mon. 11. 
MacSwain, J. W. 

1943. The primary larva and systematic position of the meloid genus Poreo- 
spasta (Coleoptera). Ann. Ent. Soc. America, vol. 36, pp. 360-64. 

1951. A new genus of Meloidae from North America. Pan-Pacific Ent., vol. 
27, p. 58. 

1956. A classification of the first instar larvae of the Meloidae (Coleoptera). 
Univ. California Publ. Ent., vol. 12. 

Maehler, K. L. 

1939. Notes on the Meloidae. Pan-Pacific Ent., vol. 15, p. 65. 
Maran, J. 

1941. De novis parumque cognitis Chinensibus generis Lytta F. speciebus 
ex affinate L. aeneiventris Haag (Coleoptera. Meloidae). Bull. Sect. Ent. 
Mus. Prague, vol. 19, pp. 180-85. 
Marshall, G. A. K. 

1902. Five years' observations and experiments (1896-1901) on the bio- 
nomics of South African insects, chiefly . . . mimicry and warning colors. 
Trans. Ent. Soc. London, 1902, pp. 287-584. 
Meyer, H. H., and R. Gottlieb. 

1926. Experimental pharmacology. Philadelphia, London, and Montreal. 
Michener, C. D. 

1944. A comparative study of the appendages of the eighth and ninth ab- 
dominal segments of insects. Ann. Ent. Soc. America, vol. 37, pp. 336-51. 

Michener, C. D., and R. R. Sokal. 

1957. A quantitative approach to a problem in classification. Evolution, vol. 
11, pp. 130-62. 

Millike n, F. B. 

1921. Results of work on blister beetles in Kansas. U.S. Dept. Agr. Bull. 967. 
Moore, J. A. 

1952. An analytical study of the geographic distribution of Rana septentrio- 
nalis. Amer. Nat., vol. 86, pp. 5-22. 

Mutchler, A. J., and H. B. Weiss. 

1924. The oil and blister beetles of New Jersey. State of New Jersey Dept. 
Agr. Circ. 76. 
Paoli, G. 

1938. Note sulla biologia e sulla filogenesi dei Meloidi. Mem. Soc. Ent. 
Italia, vol. 16, pp. 71-96. 



LITERATURE CITED IN THE TEXT 245 

Penafiel y Barranco, A. 

1866. Estudio sobre dos espeeies de cantarides mexicanas. Gaceta Med. 
Mexico, vol. 2, pp. 225-27, pis. 1 & 3. 
Peringuey, L. 

1909. Descriptive catalogue of the Coleoptera of South Africa. Family 
Meloidae. Trans. Roy. Soc. South Africa, vol. 1, pp. 165-297, pis. 22-24. 
Pritchett, A. H. 

1903. Some experiments in feeding lizards with protectively colored insects. 
Biol. Bull., vol. 5, pp. 271-87. 
Putnam, J. D. 

1876. Report on the insects collected by Captain Jones' expedition to north- 
western Wyoming in 1873. Proc. Davenport Acad. Nat. Sci., vol. 1, pp. 
187-92. 
Saunders, W. 

1876. Notes on Cantharides. Canad. Ent., vol. 8, pp. 221-28, 1 pi. 
Say, T. 

1823-24. Descriptions of coleopterous insects collected in the late expedition 
to the Rocky Mountains, performed by order of Mr. Calhoun, Secretary 
of War, under command of Major Long. Jour. Acad. Nat. Sci. Philadelphia, 
vol. 3, pp. 139-216 (1823), 238-82, 298-331, 403-62; vol. 4, pp. 88-99 
(1824). 
Selander, R. B. 

1954. Notes on the tribe Calospastini, with description of a new subgenus 
and species of Calosposta (Meloidae). Coleopterists' Bull., vol. 8, pp. 
11-18. 
1956. Geographic variation in the blister beetle Lytta biguttata (Coleoptera, 
Meloidae). Bull. Brooklyn Ent. Soc, vol. 51, pp. 116-27. 

1958. Melanism in some Meloidae (Coleoptera) from the Pinacate lava cap 
in northwestern Sonora, Mexico. Trans. Kansas Acad. Sci., vol. 61, pp. 
77-80. 

1959. The first instar larvae of some North American species of Meloidae 
(Coleoptera). Proc. Ent. Soc. Washington, vol. 61, pp. 205-13. 

1960. Restriction of the genus Lytta Fabricius (Meloidae). Coleopterists' 
Bull., vol. 14, pp. 80-86. 

Sherman, F., Jr. 

1913. The Meloidae (blister-beetles) of North Carolina (Col.). Ent. News, 
vol. 24, pp. 245-47. 
Simpson, G. G. 

1959. Anatomy and morphology: classification and evolution: 1859 and 1959. 
Proc. Amer. Philos. Soc, vol. 103, pp. 286-306. 
Skinner, H. 

1906. A new Cantharis. Ent. News, vol. 17, p. 217. 
Snow, F. H. 

1883. Lists of Lepidoptera and Coleoptera, collected in New Mexico by the 
Kansas University scientific expeditions of 1881 and 1882. Trans. Kansas 
Acad. Sci., vol. 8, pp. 35-45. 
Tanner, V. M. 

1928. The Coleoptera of Zion National Park, Utah. Ann. Ent. Soc. America, 
vol. 21, pp. 269-81. 
Tanner, V. M., and C. L. Hayward. 

1934. A biological study of the La Sal Mountains, Utah. Report No. 1 



246 BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lyttd 

(ecology). Proc. Utah Acad. Sci., Arts, and Letters, vol. 11, pp. 209-34, 

pi. 10. 
Tucker, E. S. 

1906. Determinations of some Texas Coleoptera, with records. Trans. Kansas 

Acad. Sci., vol. 20, pp. 85-89. 
Ulke, H. 

1875. Report upon the collections of Coleoptera made in portions of Nevada, 

Utah, California, Colorado, New Mexico, and Arizona during the years 

1871, 1872, 1873, 1874. In Report upon the geographical and geological 

explorations and surveys west of the 100th meridian, vol. 5, pp. 811-27, pi. 

61. Washington. 
1882. Descriptions of new species. Bull. Brooklyn Ent. Soc, vol. 4, pp. 41-42. 

[Reprint of Ulke's 1875 article.] 
1902. A list of the beetles of the District of Columbia. Proc. U.S. Nat. Mus., 

vol. 25, art. 1275. 
Van Dyke, E. C. 

1928. A reclassification of the genera of North American Meloidae (Coleop- 
tera) and a revision of the genera and species formerly placed in the tribe 
Meloini. . . . Univ. California Publ. Ent., vol. 4, pp. 395-474, pis. 15-19. 

1929. New species of Meloidae (Coleoptera). Bull. Brooklyn Ent. Soc, vol. 
24, pp. 127-33. 

1947. New species of Coleoptera from western North America. Pan-Pacific 
Ent., vol. 23, pp. 155-61. 
Vaurie, P. 

1950. The blister beetles of north central Mexico (Coleoptera, Meloidae). 
Amer. Mus. Novitates, no. 1477. 

Wellman, F. C. 

1910. The generic and subgeneric types of the Lyttidae (Meloidae s. Can- 

tharidae auctt), (Col). Canad. Ent., vol. 42, pp. 389-96. 
1912. New species of Lyttidae, with notes on described species (Coleop. ). 
Ent. News, vol. 23, pp. 29-38. 
Werner, F. G. 

1951. Additions to the nearctic Meloidae (Coleop.). Psyche, vol. 57, pp. 
131-36. 

Wickham, H. F. 

1902. A catalogue of the Coleoptera of Colorado. Bull. Lab. Nat. Hist. State 
Univ. Iowa, vol. 5, pp. 217-310. 
Wilson, E. O., and W. L. Brown, Jr. 

1953. The subspecies concept and its taxonomic application. Syst. Zool., vol. 
2, pp. 97-111. 
Wood, S. L. 

1952. Observations on the homologies of the copulatory apparatus in male 
Coleoptera. Ann. Ent. Soc. America, vol. 45, pp. 613-17. 

Xambeu, P. 

1900. Moeurs et metamorphoses des insectes. 9° memoire, deuxieme partie. 
Rev. d'Ent., vol. 19, pp. 1-56. 



Additional Figures 



Figs. 64-85. Male antennae of species of Lytta: 64, nitidicollis; 65, morrisoni; 66, 
fulvipennis; 67, lecontei; 68, variabilis; 69, scituloides (allotype); 70, michoacanae; 
71, ehenina; 72, plumbea; 73, koltzei; 74, augusti; 75, sonorae; 76, biguttata; 77, 
scitula; 78, quadrimacidata (Veracruz); 79, proteus; 80, reticidata; 81, agrestis; 82, 
mirifica; 83, deserticola; 84, tenebrosa; 85, blaisdelli (Siskiyou County, California). 



248 




249 



Figs. 86-107. Male antennae of species of Lytta: 86, melaena; 87, morosa; 88, fune- 
rea; 89, magister; 90, vulnerata; 91, peninsularis; 92, erebea; 93, eucera; 94, san- 
guinea; 95, erythrothorax; 96, mutilata; 97, cardinalis; 98, sublaevis; 99, auriculata; 
100, nigripilis (cotype); 101, hoppingi; 102, lugens; 103, aeneipennis; 104, refulgens; 
105, crotchi; 106, chloris; 107, rathvoni. 



250 




251 



Figs. 108-116. Male antennae of species of Lytta: 108, stygica (northern California); 
109, stygica (southern California); 110, moerens; 111, insperata; 112, nuttalli; 113, 
molesta; 114, polita; 115, sayi; 116, aenea. Figs. 117-129. Female antennae of species 
of Lytta: 117, nitidicollis; 118, morrisoni; 119, fulvipennis; 120, lecontei; 121, uari- 
abilis; 122, ebenina; 123, plumbea; 124, koltzei; 125, proteus; 126, augusti; 127, 
reticulata; 128, agrestis; 129, mirifica. 



252 




253 



Figs. 130-151. Female antennae of species of Lijtta: 130, deserticola; 131, tenebrosa; 
132, blaisdelli; 133, morosa; 134, funerea; 135, vulnerata; 136, eucera; 137, sanguined; 
138, erythrothorax; 139, cardinalis; 140, suhlaevis; 141, auriculata; 142, hoppingi; 
143, lugens; 144, aeneipennis; 145, refulgens; 146, chloris; 147, stygica (northern 
California); 148, moerens; 149, insperata; 150, nuttalli; 151, molesta. 



254 




255 



Figs. 152-154. Female antennae of species of Lytta: 152, polita; 153, aenea; 154, 
sayi. Figs. 155-158. Labra of species of Lytta: 155, polita; 156, aenea; 157, sayi; 
158, unguicularis. Figs. 159-167. Right elytra of species of Lytta: 159, quadrimaculata 
(showing range of variation); 160, biguttata (a, pale race; b-d, typical race; c, 
southern Mexico race); 161, scitula. Figs. 162—167. Hind trochanters of species of 
Lytta: 162, insperata, male; 163, nuttalli, male; 164, cyanipennis, male (typical); 
165, cyanipennis, male (Utah); 166, cyanipennis, female (typical); 167, cyanipennis, 
female (Utah). 



256 




J 
f 



J 




159 






257 



Figs. 168-191. Pronota of species of Lytta: 168, variabilis; 169, puberula; 170, plum- 
bea; 171, koltzei; 172, canelas; 173, biguttata; 174, proteus; 175, mirifica; 176, an- 
zonica; 177, deserticola; 178, tenebrosa 179, blaisdelli; 180, lugubris; 181, melaena; 
182, morosa; 183, magister; 184, culnerata; 185, auriculata; 186, lugens; 187, aenei- 
pennis; 188, aeneipennis (variant); 189, mittalli; 190, molesta; 191, childi. 



258 



Fig. 192. Anterior view of male middle tibia of Lytta eucera. Fig. 193. Deformed 
femur and tibia of left middle leg of female of Lytta angusti from La Parada, Oaxaca. 
Figs. 194-198. Hind tibial spurs of species of Lytta: 194, lugubris, posterior view, 
right hind leg; 195, blaisdelli, same; 196, lugens, anterior view, right hind leg; 197, 
insperata, outer spur; 198, aeneipennis, inner view, right hind leg. Figs. 199-203. 
Mesal aspect of fore tarsal claws of species of Lytta: 199, variabilis; 200, puberula; 
201, auriculata; 202, stygica; 203, moerens. Figs. 204-217. Male pygidia of species 
of Lytta: 204, moesta; 205, variabilis (variant from Temescaltepec, Mexico); 206, 
sonorae; 207, biguttata; 208, scitula; 209, quadrimaculata (a, Veracruz; b, c, Central 
Plateau); 210, proteus; 211, tenebrosa; 212, blaisdelli; 213, melaena; 214, morosa; 
215, funerea; 216, magister; 217, vulnerata. 



260 



204 



94 195 





196 






197 



198 




199 200 201 202 203 




205 



206 




216 



217 



261 



Figs. 218-221. Male fifth abdominal sterna of species of Lytta: 218, insperata; 219, 
nuttalli; 220, cyanipennis; 221, childi. Figs. 222-241. Male sixth abdominal sterna of 
species of Lytta: 222, morrisoni; 223, moesta; 224, fulvipennis; 225, variabilis; 226, 
michoacanae; 227, ebenina; 228, plumbea; 229, koltzei; 230, canelas; 231, augusti; 
232, bipuncticollis; 233, sonorae; 234, scitula; 235, quadrimaculata (Veracruz); 236, 
proteus; 237, reticulata; 238, agrestic; 239, mirifica; 240, deserticola; 241, corallifera. 



262 




218 






219 



220 



221 




222 



223 



224 



225 




226 



227 



228 



229 




230 




231 





232 



233 




234 




235 



236 




237 




238 




239 




240 




241 



263 



Figs. 242-265. Male sixth abdominal sterna of species of Lytta: 242, tenebrosa; 243, 
lugubris, 244, melaena; 245, morosa; 246, funerea; 247, magister; 248, vulnerata; 
249, peninsularis; 250, eucera; 251, sanguined; 252, erythrothorax; 253, mutilata; 254, 
sublaevis; 255, auriculata; 256, hoppingi; 257, lugens; 258, aeneipennis; 259, aenei- 
pennis (variant); 260, refulgens; 261, rathvoni; 262, chloris; 263, stygica (Argus 
Mountains, California); 264, moerens; 265, navajo. 



264 




249 




250 



251 



252 



253 






254 




255 



256 



258 




259 




260 




261 







263 



264 



265 



265 



Figs. 266-274. Male sixth abdominal sterna of species of Lytta: 266, insperata; 267, 
nigrocyanea; 268, nuttalli; 269, molesta; 270, childi; 271, polita; 272, aenea; 273, 
sayi; 274, unguicularis. Figs. 275-289. Female sixth abdominal sterna of species of 
Lytta: 275, plumbea; 276, koltzei; 277, canelas; 278, bipuncticollis; 279, biguttata; 
280, scitula (a, ventral view; b, lateral view); 281, cardinalis; 282, auriculata; 283, 
rathvoni; 284, moerens; 285, nuttalli; 286, molesta; 287, childi; 288, sayi; 289, «n- 
guicu laris. 



266 






270 



268 




269 



271 




272 



273 




286 



287 



288 



289 



267 



Figs. 290-298. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 290, nitidicollis; 291, 
morrisoni; 292, moesta; 293, fulvipennis; 294, lecontei; 295, variabilis (2 mi. S Tran- 
coso, Zacatecas); 296, variabilis (variant from Temescaltepec, Mexico); 297, sonorae 
(apex of aedeagus); 298, puberula. 



268 




290 








292 



293 




294 





297 



296 




269 



Figs. 299-306. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus ) of species of Lytta: 299, michoacanae; 300, 
ebenina; 301, plumbea; 302, koltzei; 303, canelas; 304, augusti; 305, bipuncticollis; 
306, biguttata ( typical race ) . 



270 






300 




302 





305 



306 



271 



Figs. 307-313. Male genitalia (a, ventral view of gonoforeeps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 307, scitula; 308, 
quadrimaculata (a, b, c, Veracruz; c', typical specimen from Central Plateau; c", 
variant from Central Plateau); 309, proteus; 310, reticulata; 311, agrestis; 312, miri- 
fica; 313, cribrata. 



272 





307 




309 










273 



Figs. 314-321. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 314, deserticola^ 315, 
corallifera; 316, tenebrosa; 317, blaisdelli ( Siskiyovi County, California); 318, lugu- 
bris; 319, melaena; 320, morosa; 321, funerea. 



274 




314 









320 



321 



275 



Figs. 322-329. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 322, magister; 323, 
vulnerata; 324, peninsalaris; 325, erebea; 326, eticera (c, variant from Finca Gibral- 
ter, Mexico (state unknown); c", variant from Tancitaro, Michoacan); 327, san- 
guined; 328, erythrothorax; 329, mutilata. 



276 






322 



323 






324 



325 







326 



327 








328 



329 



277 



Figs. 330-338. Male genitalia (a, ventral view of gonoforeeps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 330, cardinalis; 331, 
sublaevis; 332, auriculata (c', variant); 333, nigripilis (cotype) (c', variant); 334, 
hoppingi; 335, lugens; 336, aeneipennis; 337, refulgens; 338, crotchi. 



278 





330 









b r c 



332 



333 






334 



335 




336 





338 



Figs. 339-346. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 339, chloris (Kern 
County, California); 340, stygica (Mendocino County, California); 341, moerens; 
342, navajo; 343, insperata; 344, nigrocyanea; 345, nuttalli; 346, molesta. 



280 





b c 




339 



340 







a v b £s c a 

341 342 




b 6/ c 








343 



344 








345 



346 



281 



Figs. 347-350. Male genitalia (a, ventral view of gonoforceps; b, lateral view of 
gonoforceps; c, lateral view of aedeagus) of species of Lytta: 347, childi; 348, 
polita; 349, aenea; 350, sayi. 



282 






347 



348 







349 




350 



283 



Inde 



x 



Specific scientific names are indexed under appropriate generic names, except 
in the case of species of the genus Lytta. Junior synonyms are printed in 
italics; homonyms are followed by authors' names. For taxa of the genus 
Lytta, page numbers in italics indicate principal references. 



abbreviata, Lytta, 24 

Abdominal sterna: terminology of, 5 

Acrolytta, 24 

Activity period, adult, 11 

Adicolytta: 23, 26, 27, 29, 30, 32, 
137-140; feeding habits, 13; modi- 
fications of males, 14; distribution, 
32; phylogeny, 138-140 

adonis, Lytta, 24 

aenea, Lytta: 35, 36, 225, 229-234; 
attacking fruit trees, 9; invalidly 
designated type of Pomphopoea, 
223; Texas race, 234; of Say, 235 

Aenea Group, 27, 226, 228-229 

aeneipennis, Lytta, 3, 39, 162, 174- 
175 

aeneiventris, Lytta, 25, 27 

Aeneiventris Group, 27 

Afrolytta: 24, 25; amoena, 25 

agrestis, Lytta, 3, 37, 102, 105-106, 
108 

Aguascalientes, 95 

Alabama, 227, 230, 239 

Alberta, 207, 213 

alemani, Lytta, 87, 90 

Alfalfa, 61, 105, 213 

alfredi, Lytta, 87 

Amelanchier, 234 

amoena, Lytta. See Afrolytta 

Amsinckia, 165 

Andrena, 18 

angidcollis, Lytta, 131, 132, 136 

Anomalies: of head, 60; of legs, 82; 
of pronotum, 135-136 

Antennae: sexual dimorphism of, 3-4; 
method of measuring, 5 

antennalis, Lytta, 27 

Anthophora: linsleyi, 18, 118, 185, 
192; stanfordiana, 200 

Anthophoridae, 17 



Ants, 10 

apicalis, Lytta, 25, 28 

Apis mellifera, 18 

Aposematic coloration. See Coloration 

Apples, 9, 234 

Applopappus spinulosus, 90 

Arctostaphylos, 192 

Arctotis, 192 

arborea, Lytta, 185, 186, 191-192 

Argemone: 97, 169, 174; platyceras, 

60, 73, 76, 96, 147 
Arizona, 58, 69, 88, 106, 108, 111, 

112, 123, 129, 133, 153, 167, 188, 

201, 208, 214 
arizonica, Lytta, 37, 103, 107, 108 
Artemesia, 136 
Asclepias, 67 
Astragalus, 192, 201, 209 
atrovirens, Lytta, 77, 78, 79 
augusti, Lytta, 42, 51, 74, 80-82 
auriculata, Lytta: 3, 43, 45, 46, 162, 

165-169; allopatric relationship 

with nigripilis, 170-171 
azalea, 239 

Baboons: refuse Meloidae, 11 

Baccharis, 136 

badakschanica, Lytta, 26 

Baeria: 118, 185; chrysostoma, 185 

Bahia: absinthifolia, 90; pedata, 90 

Baileya multiradiata, 90 

Baja California Norte, 55, 134, 154, 

168, 219 
Baja California Sur, 123, 136, 142, 

219 
Barley, 9, 209 
bayoni, Lytta, 25 

Beans: 9, 61, 209, 212, 238; lima, 200 
Bees: 1, 8, 13, 19; species attacked, 

17-18; laboratory learings with pol- 



285 



286 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



len of, 18-19. See also specific fam- 
ilies and genera 

Beets: 19, 202, 209; sugar, 105 

benguellana, Lytta, 25 

Bering Strait, 224 

Bidens leucantha, 97 

bieti, Lytta, 26 

bifasciata, Lytta, 93 

bifasciatus, Lytta, 93, 94, 95 

biguttata, Lytta: 3, 13, 38, 51, 85, 
86-90; distribution, 35; typical race, 
89, 90; pale race, 89; southern 
Mexico race, 89, 90 

bilineatipennis, Lytta, 24 

binotatithorax, Lytta. See Acrolytta 

bipartite, Lytta, 97, 101 

bipuncticollis, Lytta, 40, 51, 74, 82- 
84 

Birds: predation by, 11, 213; falcon, 
11; magpie, 213 

bivirgata, Lytta, 86, 88, 90 

blaisdelli, Lytta, 45, 49, 114, 116, 
119-120, 200 

Blister beetles, 1. See also Meloidae 

Boraginaceae, 50, 130, 165, 193 

British Columbia, 133, 211, 213 

brucci, Lytta, 25 

bruchi, Lytta. See Acrolytta 

Bur, creeping, 105 

Butterflies, 220 

Butternut, 238 

California: 32, 55, 56, 57, 117, 119, 
121, 123, 126, 129, 133, 153, 164, 
167, 170, 172, 174, 175, 177, 178, 
179, 180, 188, 192, 197, 202, 208, 
211, 217, 219; number of species 
in, 32 

Camouflage, 97 

Campanula, 70 

Campanulaceae, 50, 62, 70 

Canada, 31, 49, 133, 157, 205, 207, 
209, 211, 213, 223, 235, 236, 237, 
238 

canelas, Lytta, 41, 74, 79-80 

Cantharides: preparation of, 9; 
sources of, 9; plaster, 10 

Cantharidin, 9-10 

Cantharis, 21, 22, 25 

Capparidaceae, 90 

Caprifoliaceae, 13, 31, 169, 238 



Caragana: 13, 209, 215; arborescens, 
209 

caraganae, Lytta: 27; food plants, 13 

cardinalis, Lytta: 38, 140, 148, 155- 
156; of Horn, 3, 58 

cardonii, Lytta, 26 

carneola, Lytta, 25 

Carya, 234 

Castilleja, 120 

catamarcensis, Lytta. See Afrolytta 

Ceanothus: 161, 171, 177, 178, 192; 
cuneatus, 193 

Central Plateau of Mexico, 32, 64, 75, 
84, 94, 113 

Central Valley of California, 118, 134, 
164, 165, 170, 172, 186, 189, 192, 
198, 199, 217, 218, 220, 222 

Centromadia, 127 

Ceratina, 2, 18 

Cercocarpus, 192 

Cercopithecus monkey: refuses Mel- 
oidae, 11 

Ceylon, 31 

Character convergence. See Conver- 
gent evolution 

Cherry, 238 

Chiapas, 145 

Chihuahua, 64, 89, 95, 110, 154 

childi, Lytta, 3, 41, 45, 46, 216, 218- 
222 

Childi Subgroup: 26, 160, 194, 215- 
216; phylogeny, 160 

China, 31 

chloris, Lytta: 2, 3, 46, 48, 163, 179- 
185, 192; larval host, 18; Coastal 
Range race, 182, 183; Sierran foot- 
hills race, 182-183 

Chrysothamnus: 136; paniculatus, 
130; nauseosus, 136; stenophyllus, 
136 

clematidis, Lytta, 157, 158 

Clematidis Group, 26, 158 

Cleome serrulata, 90 

Climatic variation: effect on develop- 
ment, 12 

Clover, 147 

Cnemidophacos pectinatus, 209, 215 

Coahuila, 68, 88, 145, 154 

Coarctate larval instar. See Larvae 

Coastal Range, 125, 164, 169, 170, 
180, 181, 182, 183, 186, 189, 190, 
192, 193 



INDEX 



287 



coccinea, Lytta, 28 

Coldenia palmed, 130 

Colima, 98, 145, 152 

Colletes: 17; fulgidus, 200 

Colletidae, 17 

colon, Lytta. See Acrolytta 

Colorado, 58, 69, 88, 103, 186, 203, 
208, 214 

Colorado Desert, 186 

Coloration: aposematic, 137-138; 
structural, 189 

comans, Lytta, 46, 48, 163, 192-193; 
compared with chloris, 182, 183, 
185 

Competition, 223 

Compositae, 13, 30, 31, 50, 60, 61, 
90, 93, 96, 115, 118, 122, 127, 130, 
136, 147, 161, 165, 169, 185, 192, 
213, 222 

compressicornis, Lytta. See Linsleya 

Connecticut, 230, 236, 239 

Convergent evolution: within Poreo- 
spasta, 31, 159-160; within Para- 
lytta, 51; of tarsi, 51, 117; in 
scitula and scituloides, 68; of head 
modifications, 68; in Lytta and 
Oenas, 92; in bipuncticollis and 
proteus, 99; of male genitalia, 99, 
160; in tenebrosa and viridana, 117; 
of antennae, 117, 159-160; within 
Mutilata Group, 140; in Paralytta 
and Poreospasta, 159; in Childi and 
Cyanipennis subgroups, 160; char- 
acter convergence, 220-222 

convexa, Lytta. See Linsleya 

Convolvulaceae, 13, 30, 32, 50, 62, 
67, 72, 73, 79, 102, 105, 138, 150, 
155, 156, 169 

Convolvulus arborea, 67 

cooperi, Lytta, 3, 131, 132, 135, 136 

corallifera, Lytta, 37, 51, 109, 112- 
113 

Cornaceae, 238 

Cornus, 238 

Costa Rica, 146 

Cotton, 61 

Courtship, 14 

Cowania stansburiana, 192 

Crataegus, 234 

cribrata, Lytta, 3, 37, 109-111 

crotchi, Lytta, 3, 41, 162, 177-178 

Cruciferae, 192, 200 



Cryptantha, 193 

Cucujoidea, 1 

Cucurbita moschata, 147 

Cucurbitaceae, 30, 138, 147 

Cusculata umbellata, 169 

Cuscuta, 13, 155 

cyanescens, Lytta, 76, 78, 79 

cyanipennis, Lytta: 3, 13, 32, 44, 47, 
205, 209-213; swarming, 1; attack- 
ing field crops, 9; seasonal distribu- 
tion, 11; behavior, 12, 19; feeding 
habits, 13; courtship, 14; mating, 
15; oviposition, 15-16; incubation, 
16; number of larval instars, 17; 
reared in laboratory, 18-19; first 
instar larva, 19; grub stage, 19 

Cyanipennis Subgroup: 26, 160, 194, 
204-205; distribution, 35; phylo- 
geny, 160 

Daisy, African, 169, 192 

Death feigning, 12 

deserti, Lytta, 28 

deserticola, Lytta, 3, 37, 109, 111- 
112 

Deserticola Subgroup, 26, 102, 108- 
109 

Diadasia bituberculata, 18, 123 

Dicentra, 169 

dichroa Fischer von Waldheim, Lvtta, 
26 

dichroa LeConte, Lytta, 3, 60, 61 

difficilis, Lytta, 3, 185, 186, 191 

Diholcos bisulcatus, 209, 215 

dimidiata, Lytta. See Acrolytta 

discipennis, Lytta, 26 

Dispersal: barriers or deterrents to, 
35, 223. See also Distribution, geo- 
graphic 

Distribution, geographic: of Meloidae, 
1; of Lytta, 31-32, 35; centers of, 
32; of Poreospasta, 32, 157; effect 
of glaciation on, 35; of Paralvtta, 
49-50; of Variabilis Group, 51; of 
Reticulata Group, 51, 102; of Ma- 
gister Group, 51, 114; of Nitidi- 
collis Group, 53; of Fulvipennis 
Group, 58; disjunct, 70, 206, 223- 
224, 233-234, 237; of Koltzei Sub- 
group, 73; Quadrimaculata Sub- 
group, 84; of Tenebrosa Subgroup, 
115; of Magister Subgroup, 127; 



288 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



of Adicolytta, 137; of Stygicia 
Group, 161; allopatric, 170-171, 
193, 214, 223, 239; sympatric, 182, 
220; relict, 187; of Moerens Group, 
194; of Moerens Subgroup, 195; of 
Cyanipennis Subgroup, 205; of 
Childi Subgroup, 215; of Pompho- 
poea, 223; of Sayi Group, 235. See 
also Dispersal 

Distribution, seasonal, 11-12 

District of Columbia, 230 

Distrito Federal, 76, 78, 95, 98, 145 

dolosa, Lytta, 185, 186, 191 

Durango, 80, 88, 95, 110, 146, 154 

Eastern Deciduous Forest, 35, 223 

ebenina, Lytta: 12, 41, 42, 51, 63, 
72-73; behavior, 12 

Ecological relationships: of crotchi 
and aeneipennis, 178; of childi and 
molesta, 220 

Economic importance: damage to cul- 
tivated plants, 9; source of canthar- 
idin, 9 

Eggs: contain cantharidin, 91; incu- 
bation of, 16; number of, 16; of 
quadrimaculata, 97. See also Op- 
position 

Elder, 238 

elegantula, Lytta, 25 

Eletica, 11 

Emergence, adult, 12 

Encelia, 169 

encera, Lytta, 144 

England, 31 

enona, Lytta, 25 

Epicauta: 12, 16, 121; source of can- 
tharidin, 9; eaten by vertebrates, 
11; number of larval instars, 17; 
lugubris Haag-Rutenberg, 121 

Epicautini, 8, 23 

erebea, Lytta: 43, 138, 139, 143-144; 
distribution, 32 

Ericaceae, 192, 239 

Eriodictyon: 169, 193; calif ornicum, 
192 

Eriogonum, 171, 174 

Erodium cicutarium, 217 

erythrothorax, Lytta, 40, 60, 140, 148, 

150-152 
erytrotora, Lytta, 150 
erytrotorax, Lytta, 150 



Eschscholtzia: 97, 161, 165, 169, 175, 
178, 179, 192; californica, 192, 193 

Ethiopian region: fauna of, 25 

Eucera, 18 

eucera, Lytta: 3, 13, 32, 43, 137, 139, 
143, 144-147; seasonal distribution, 
11; behavior, 12; feeding habits, 13 

Eucera Group: 26, 140, 143; phylo- 
geny, 138-139 

Eupomphini, 23, 52, 53 

Eurasian fauna, 25-26 

exclamans, Lytta, 25 

Fagaceae, 234 

fairmairei, Lytta, 157 

Fairmairei Group, 26, 157-158 

Falcon: refuses Meloidae, 11 

Fall, H. C: classification of, 3-4 

fasciolata, Lytta, 93, 94, 96 

Feeding habits: of Meloidae, 8; ag- 
gregations, 9; summary, 13-14; asso- 
ciations, 76, 97; of plumbea, 76; 
of quadrimaculata, 97; of tene- 
brosa, 119; of cyanipennis, 212- 
213. See also Food plants 

femoralis, Lytta, 227, 228 

Filaree, 217 

filiformis, Lytta, 229, 230, 234 

fissiceps, Lytta, 24 

fissicollis, Lytta, 25, 26 

flava, Lytta, 26 

flavicollis, Lytta, 26 

flavipennis, Lytta, 27 

naviventris, Lytta, 26 

navoangulata, Lytta, 26 

Flies, ceratapogonid: attacking Mel- 
oidae, 10 

Florida, 227 

Food plants: records lacking for many 
species, 8; cultivated, 9; of most 
importance, 13; specificity, 13; of 
Paralytta, 30, 50; of Adicolytta, 30; 
of Lytta (subgenus), 31; of Poreo- 
spasta, 31, 157; of Pomphopoea, 
31; and competition, 32; of Vari- 
abilis Subgroup, 62; of Koltzei Sub- 
group, 73; of Quadrimaculata Sub- 
group, 84; of Stygica Group, 161. 
See also Feeding habits and spe- 
cific families and genera of plants 

France, 78 

Fraxinus, 13 



INDEX 



289 



Frogs. See Rana 

fryi, Lytta, 25 

fulgifer, Lytta, 205 

fulgifera, Lytta, 206 

fulviceps, Lytta, 26 

fulvipennis, Lytta: 3, 38, 52, 58-60; 

male antennae, 3-4; not eaten by 

lizards, 11 
Fulvipennis Group, 26, 32, 52, 53, 

57-58 
Fumariaceae, 169 
funerea, Lytta, 3, 43, 51, 52, 114, 

122, 126-127 

Generations, 12 

Genitalia: male, terminology of, 5; 
adaptations for coupling, 15 

gentilis, Lytta. See Linsleya 

Geographic distribution. See Distri- 
bution, geographic 

Geographic variation: treatment of, 
4-5; polymorphic, in variabilis, 65- 
67; in augusti, 82; in quadrimacu- 
lata, 96; in tenebrosa, 117-118; in 
blaisdelli, 119-120; melanistic, in 
magister, 130; in vulnerata, 134- 
135; in sublaevis, 165; in auricu- 
lata, 168-169; in stygica, 189-191; 
in moerens, 198-199; in cyanipen- 
nis, 211-212; in childi, 219-220; 
character or sympatric convergence 
in, 220-222; centripetal cline, 220; 
in aenea, 231-234; in sayi, 237-238 

Georgia, 227, 230 

Geraea canescens, 130, 169 

Geraneaceae, 217 

Gerrhonotus lizards: reaction to fulvi- 
pennis, 11 

glabricollis, Lytta, 26 

Gramineae, 61, 238 

Grass, 152, 165. See also Gramineae 

Grasshoppers: larval hosts of Meloi- 
dae, 1, 8 

Great Basin, 32, 35, 132, 135, 186 

Great Plains, 35, 51, 58, 88, 103, 223 

Gregariousness. See Swarming be- 
havior 

griseopubescens, Lytta. See Acrolytta 

Grub stage. See Larvae 

Guanajuato, 64, 76, 89, 95, 99, 146, 
152 

Guatemala, 146 



Guayule, 90 

Guerrero, 71, 83, 91, 99, 149, 152, 

156 
Gutierrezia, 136 

Hadronema militaris: attacking nut- 

talli, 11, 209 
Halictus, 18 
Helianthus, 60 
Heliconids, 220 
Hemiptera, 10. See also Hadronema 

militaris 
Hemizonia, 127 
Hidalgo, 65, 95, 113, 146 
Holarctic region, 35 
Honey bee. See Apis mellifera 
hoppingi, Lytta, 39, 40, 162, 171-172 
Horiini, 9 

Horn, G. H.: classification of, 3-4 
Hornia, 8 

humilis, Lytta, 77, 78, 79 
Hybridization: 209, 222; of viridana 

and nuttalli, 215 
Hydrophyllaceae, 169, 192, 193 
Hymenoptera: larval hosts of Meloi- 

dae, 2. See also Bees 
Hymenoxys richardsoni, 90 
Hypermetamorphosis, 1, 8 

Idaho, 119, 133, 189, 208, 211 

Illinois, 231, 236, 239 

impressicollis, Lytta, 26 

incommoda, Lytta, 3, 196, 199 

Incubation. See Eggs 

India, 31 

Indiana, 231 

Indiolytta, 27, 28, 29, 30, 31 

inflaticeps, Lytta, 24 

Insecticide: cantharidin as an, 10 

insperata, Lytta, 3, 44, 47, 160, 195, 
201-202 

insperatus, Lytta, 201 

intricata, Lytta, 109, 110-111 

Introgression, 183 

Iowa, 236 

Ipomoea: 67, 150; longifolia, 67; 
pringlei, 67; stans, 67; leptophylla, 
105. See also Morning-glory 

Iridaceae, 192 

Iris, 192 

Italy, 9 



290 



BIONOMICS, SYSTEMATICA, AND PHYLOGENY OF Lytta 



Jalisco, 65, 95, 99, 146, 152, 156 
Japan, 31 
Juglandaceae, 234, 238 

Kalmia, 239 

Kansas, 59, 88, 104, 206, 213 

Kentucky, 231 

Kolkwitzia, 238 

koltzei, Lytta: 42, 74, 77-79; of 

Champion, 74 
Koltzei Subgroup, 26, 51, 62, 73-74 
kwanhsiensis, Lytta, 27 

Labiatae, 30, 50, 55, 61, 169, 192 

laeta, Lytta, 26 

Lampropeltis, 220 

Larvae: mode of life in Meloidae, 1, 
2; knowledge of development of, 
8; number of instars, 8, 16-17; 
adaptive significance of burrowing 
habit of, 11; nomenclature of in- 
stars, 17; coarctate instar, 8, 16, 19- 
20; grub stage, 17, 19; ultimate in- 
star, 20. See also Larvae, first instar 

Larvae, first instar: 19, 27; phoresy 
by, 1; first described, 2; contain 
cantharidin, 9; of Magister Group, 
51-52, 114; of Moerens Group, 158; 
of Poreospasta, 158, 160; of Stygica 
Group, 158. See also Larvae and 
Larval hosts 

Larval hosts: 1, 12, 13; summary of, 
17-18; of vesicatoria, 18; number 
of cells required, 18; of variabilis, 
67; of tenebrosa, 118-119; of me- 
laena, 123-124; of chloris, 185, 
192; of stygica, 192; of moerens, 
200. See also Larvae and Larvae, 
first instar 

Lathyrus, 212, 215 

Laurel, mountain, 239 

Layia, 185, 192 

lecontei, Lytta, 3, 38, 52, 58, 60-61 

Leguminosae, 13, 31, 50, 61, 90, 130, 
147, 161, 169, 171, 174, 177, 179, 
192, 194, 200, 201, 209, 212, 215, 
217, 222, 238 

Lesquerella gordani, 192 

Lettuce, 213 

Ligustrum, 13 

Liliaceae, 50, 130 

Linsleya: 3, 22; gentilis, 3; compres- 



sicornis, 3, 22; suavissima, 3, 22; 

convexa, 22; sphaericollis, 22, 202, 

213 
Lizards, 11 
Longevity, adult, 12 
Lonicera, 73 
Louisiana, 227, 231 
lucida, Lytta. See Prolytta 
lugens, Lytta, 3, 41, 45, 46, 162, 172- 

174 
lugubris Horn, Lytta, 3, 45, 49, 116, 

120-121 
lugubris Ulke, Lytta, 120, 121 
Lupine. See Lupinus 
Lupinus, 130, 169, 171, 174, 177, 

179, 192, 200, 212, 217, 222 
luteovittata, Lytta, 157 
Luteovittata Group, 26, 157 
Lycium, 192 
Lytta (subgenus): 27, 29, 30, 31; 

feeding habits, 13; food plants, 13, 

31; distribution, 31 
Lyttinae, 22 
Lyttini: 1, 22, 23, 24, 25, 53; South 

American, 24 

maculicollis, Lytta, 218, 222 
magister, Lytta: 3, 36, 51, 114, 128- 

130; adult activity period, 11; type 

of Paralytta, 49 
Magister Group: 26, 52, 53, 102, 113- 

115; of MacSwain, 4; distribution, 

32, 51; phylogeny, 51; first instar 

larvae, 51-52 
Magister Subgroup, 26, 114, 127-128 
Magpie: predation by, 213 
malatiensis, Lytta, 26 
Malus, 234 
Malvaceae, 61, 169 
Mammals: predation by, 11 
Manchuria, 31 
manicata, Lytta, 26 
Manitoba, 213 
margarita, Lytta, 3, 37, 114, 128, 236- 

137 
marginicollis, Lytta, 26 
Maryland, 231 
mashuna, Lytta. See Prolytta 
Massachusetts, 231, 236 
Mating: 15, 97; position during, 14; 

adaptations of genitalia for, 15; 

interspecific, 169 



INDEX 



291 



Megachile, 18 

melaena, Lytta: 2, 3, 44, 49, 122-124; 
larval host, 18 

Melaena Subgroup, 26, 115, 122 

Melanism: in magister, 130 

melanura, Lytta, 26 

Meloe: 8, 22; compared with Lytta, 
24; resemblance to sublaevis, 165 

Meloidae: distribution, 1; hypermeta- 
morphosis in, 1; larvae and larval 
life, 1, 8; parasitism by, 1; phoresy 
by, 1; vernacular name, 1; feeding 
habits of adults, 8; odor of, 9; as 
source of cantharidin, 9 

Meloinae, 8, 22, 31; oviposition, 8; 
mating position, 14 

menetriesi, Lytta, 27 

Menetriesi Group, 27 

Mesolytta, 27, 28, 29, 30; distribu- 
tion, 31 

Mexico (state), 65, 78, 89, 95, 99, 
144, 146 

Michoacan, 71, 76, 78, 96, 99, 144, 
146, 149, 152 

michoacanae, Lytta, 12, 40, 63, 70-71 

Mimicry, 220-222 

Minnesota, 208, 236 

minor, Lytta, 77 

Miridae, 209 

mirifica, Lytta, 37, 102, 107 

Mississippi, 227 

Missouri, 231 

moerens, Lytta, 2, 3, 9, 44, 45, 48, 
157, 160, 195, 196-200, 202 

Moerens Group: 26, 32, 161, 193- 
194; of MacSwain, 4; food plants, 
31; distribution, 32; larvae, 158; 
phylogeny, 158-160 

Moerens Subgroup: 26, 194-196; phy- 
logeny, 160 

moesta, Lytta, 22, 37, 52, 54, 56-57, 
127 

Mojave Desert, 35, 118, 132, 134, 
135, 186 

molesta, Lytta, 3, 41, 45, 47, 215, 
216-217 

Monarda citridosa, 61 

monilicornis, Lytta, 74, 76 

Montana, 88, 208, 211 

Morelos, 99, 146, 149, 152, 156 

Morning-glory, 67, 79, 105. See also 
Ipomoea 



morosa, Lytta, 3, 22, 44, 49, 114, 122, 

124-126 
morrisoni, Lytta, 37, 52, 54, 56 
Mouche d'Espagne, 9 
muelleri, Lytta, 24 
mutilata, Lytta: 3, 13, 40, 137, 140, 

148, 152-155; feeding habits, 13; 

distribution, 32; type of Adicolytta, 

137 
Mutilata Group: 26, 140, 147-148; 

phylogeny, 138, 139-140 
Mylabrini: larval food, 8; eaten by 

vertebrates, 11 
Mylabris: source of cantharidin, 9; 

number of larval instars, 17 
myrmido, Lytta, 25 

namagua, Lytta, 25 

navajo, Lvtta, 45, 46, 160, 195, 200- 

201 
Nayarit, 99, 152 
Nebraska, 59, 88, 104, 208 
neivai, Lytta, 24 

Nematodes: attacking Meloidae, 10 
Nemognatha: distribution compared 

with that of Lytta, 35 
Nemognathinae, 8 
Nevada, 119, 121, 130, 133, 189, 198, 

211 
nevadensis, Lytta, 185, 191 
New Hampshire, 231 
New Jersey, 231, 236 
New Mexico, 69, 88, 89, 104, 107, 

112, 153, 208, 214 
New York, 231, 236 
nigricornis, Lytta, 229, 230, 234 
nigrifinis, Lytta, 25, 26 
nigripilis, Lytta, 3, 45, 46, 162, 169- 

171 
nigrocyanea, Lytta, 32, 44, 46, 160, 

196, 202-204 
nigropicta, Lytta. See Acrolytta 
nitidicollis, Lytta, 3, 38, 39 52, 54- 

55 
Nitidicollis Group: 26, 32, 52, 53-54, 

127; of MacSwain, 4 
nitidula, Lytta. See Prolytta 
Nomia melanderi, 19 
North American fauna, 22 
North Dakota, 208, 227, 231, 239 
Nuevo Leon, 96 
nunemacheri, Lytta, 196, 199-200 



292 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



nutalli, Lytta. See nuttalli, Lytta 
nuttali, Lytta. See nuttalli, Lytta 
nuttalli, Lytta: 3, 32, 44, 47, 205-209; 
attacking field crops, 9; preyed on 
by mirid bug, 11; seasonal distri- 
bution, 11 

Oak, 234 
Oats, 9, 61, 209 

Oaxaca, 81, 83, 91, 99, 146, 156 
occipitalis, Lytta, 2, 3, 4, 116, 118 
octomaculata, Lytta, 93, 94, 96 
Oenas, 92 
Ohio, 231 

Oklahoma, 59, 104, 231 
Old World: subgenera of, 27-28 
Olea, 13 

Oleaceae, 13, 31 
Ontario, 207, 236 
Operculina ornithopoda, 150 
Oregon, 119, 121, 133, 189, 198, 211 
Oriental region: fauna of, 25 
Orthocarpus, 192 
Osmia, 18 

Oviposition: 15-16; in Meloinae, 8; 
in Nemognathinae, 8. See also Eggs 

Palearctic region: 157; fauna of, 25 

pollens, Lytta, 87, 88, 90 

pallidipennis, Lytta. See Prolytta 

Panama, 31, 73, 78, 79, 137, 145, 146 

Paniculolytta: 23; sanguineoguttata, 
22, 23 

Papaveraceae, 13, 30, 31, 50, 60, 62, 
73, 96, 147, 161, 165, 169, 174, 
175, 178, 179, 192, 193 

Paralytta: 15, 23, 26, 27, 29, 30, 49- 
52, 159; feeding habits, 13-14; 
modifications of males, 14; distri- 
bution, 32; phylogeny, 50-52 

Parasitism: 1, 8, 13; extent of, 17-18 

Peaches, 9, 234, 238, 239 

Pears, 9, 234, 238 

Peas, 9, 212 

Peavine, 212, 215 

pedestris, Lytta, 227 

peninsularis, Lytta, 3, 32, 39, 138, 
141-142 

Peninsularis Group, 26, 140, 141 

Pennsylvania, 231, 237 

Pentstemon: 120; spectabilis, 169 

Petalostemum, 90 



Phacelia, 169, 192 

philippi, Lytta, 24 

Phoresy, 1 

Phylogeny: principles followed in, 4; 
theoretical basis of systematics, 4; 
static relationships and, 4, 158; of 
subgenera, 30-31; of Paralytta, 50- 
52; of Reticulata Group, 50-51; of 
Variabilis Group, 50-51; of Magis- 
ter Group, 51-52, 114; of Eucera 
Group, 138-139; of Adicolytta, 138- 
140; of Mutilata Group, 139-140; 
of Poreospasta, 157-160; of Stygica 
Group, 158-159; of Moerens Group, 
159-160; of Childi Subgroup, 160; 
of Cyanipennis Subgroup, 160; of 
Moerens Subgroup, 160; of Pom- 
phoea, 224-225 

Pica pica hudsonica, predation by, 
213 

Picnoseus, 24 

pilsbryi, Lytta, 11, 109 

Pirophorum, 234, 238 

Place names: spelling of, 5 

Pleistocene: effect of glaciation on 
distribution, 35, 234; pluvial period 
in, 187; post-Pleistocene climate, 
223 

plumbea, Lytta: 13, 42, 43, 74-76; 
of Champion, 79, 80; of Vaurie, 79 

Plums, 9, 234, 238 

Pluvial Period, 187 

poeciloptera, Lytta, 157, 158 

Polita Group: 26, 225, 226; of Mac- 
Swain, 4, 158 

polita Horn, Lytta, 22, 157, 163, 165 

polita Say, Lytta: 36, 163, 225, 226- 
228; attacking fruit trees, 9; sea- 
sonal distribution, 11 

Polygonaceae, 171, 174 

Polymorphism. See Variation 

Pomphopoea: 3, 9, 21, 22, 23, 26, 27, 

28, 30, 31, 222-226; feeding habits, 
13; food plants, 31; distribution, 
35; phylogeny, 224-225 

Poppy, 76, 97 

Populus, 13 

Poreospasta: 11, 21, 22, 23, 26, 27, 

29, 31, 32, 157-161; feeding hab- 
its, 13; modifications of males, 14; 
distribution, 31; phylogeny, 157- 
160 



INDEX 



293 



Predation: on Meloidae, 10-11; by 

vertebrates, 11; by birds, 11, 213; 

by Hadronema militaris, 209 
Primrose, 61 
Primulaceae, 50, 61 
Prolytta, 25 
Prolyttini, 25 
proteus, Lytta, 38, 39, 40, 43, 85, 97- 

101 
Prunus, 192, 234, 238 
Pseudolytta, 14, 24, 27, 29, 30, 31 
Ptilothrix sumichrasti, 18, 67 
puberula, Lytta, 3, 41, 42, 51, 62, 63, 

68-70 
Puebla, 72, 78, 91, 96, 99, 146, 156 
punctata, Lytta, 26 
Pupal stage, 20 
purpurescens, Lytta, 2, 3, 185, 186, 

191 
pyrivora, Lytta, 235, 236, 238 
Pyrota, 12, 23, 24 

quadrilineata, Lytta. See Acrolytta 
quadrimaculata, Lytta: 3, 13, 38, 40, 

85, 93-97; behavior, 12; feeding 

habits, 13 
Quadrimaculata Subgroup, 26, 51, 62, 

84-85 
Quebec, 236 
Queretaro, 65, 96 

Races: vernacular names for, 4-5; of 
biguttata, 89; of tenebrosa, 117- 
118; of vulnerata, 135; of chloris, 
182; of aenea, 234. See also Geo- 
graphic variation 

Radishes, 9, 200 

Rafinesquia neomexicana, 192 

Rana: clamitans, 220; septentrionalis, 
220 

Range. See Distribution, geographic 

Ranunculaceae, 171 

rathvoni, Lytta: 3, 45, 47, 162, 178- 
179; compared with chloris, 182, 
183 

Reflex bleeding, 2 

refulgens, Lytta, 3, 41, 162, 175-177 

regis-zahiri, Lytta, 27 

Relicts, 187 

reticulata, Lytta: 3, 37, 51, 102, 103- 
105; distribution, 35 



Reticulata Group: 15, 26, 32, 51, 52, 
53, 101-102; phylogeny, 50-51 

Reticulata Subgroup, 26, 51, 102-103 

Rhamnaceae, 161, 171, 177, 178, 192, 
193 

Rhododendron, 239 

Robinia pseudo-acacia, 238 

roborowski, Lytta, 27 

Rocky Mountains, 32, 35, 127, 159, 
205 

Rosa, 238 

Rosaceae, 13, 192, 234, 238, 239 

Rose, 239 

rubra, Lytta, 26 

rubrolineata, Lytta, 25 

Sage, 118 

Salicaceae, 13, 31, 234, 238 

salicis, Lytta, 209, 210, 212 

Salix: 13; discolor, 238 

Salvia: 169, 192; multiflora, 55 

Sambucus, 13, 169 

sanguinea, Lytta: 13, 38, 137, 138, 
140, 148-150; behavior, 12; feeding 
habits, 13 

sanguineoguttata, Lytta. See Panicu- 
lolytta 

San Luis Potosi, 65, 88, 96, 113, 146 

Saskatchewan, 207, 213 

sayi, Lytta: 36, 225, 235-238; attack- 
ing fruit trees, 9; type of Pompho- 
poea, 223 

Sayi Group, 27, 226, 234-235 

Sceloporus lizards, 11 

scitula, Lytta, 13, 38, 40, 43, 85, 90- 
93 

scituloides, Lytta, 11, 40, 63, 67-68 

Scrophulariaceae, 115 

Scymus, 97 

Seasonal distribution. See Distribu- 
tion, seasonal 

Sechium edule, 147 

semilineata, Lytta. See Prolytta 

seminigra, Lytta, 24 

Sicyos angulata, 147 

Sierra Nevada, 32, 35, 115, 119, 170, 
180, 186, 187, 198, 206 

sifanica, Lytta, 27 

signaticollis, Lytta, 90, 93 

Sinaloa, 152 

smaragdina, Lytta, 185 

smaragdula, Lytta, 185, 186, 191 



294 



BIONOMICS, SYSTEMATICS, AND PHYLOGENY OF Lytta 



Snake, coral, 220 

sobrina, Lytta, 97, 101 

Solanaceae, 192 

Solidago, 136 

Sonora, 86, 112, 123, 130, 152 

sonorae, Lytta, 38, 51, 85-86 

Sonoran Desert, 32, 35, 86, 167 

South American fauna, 24 

South Carolina, 227, 231 

South Dakota, 88, 208 

Spain, 31 

Spanish fly, 9 

Speciation, 158 

Species: number of, 1 

spectabilis, Lytta, 144, 145, 147 

Sphaeralcea: 169; ambigua, 169 

sphaericollis, Lytta. See Linsleya 

Spiders: predation by, 10 

spilotella, Lytta, 25 

Spilotella Group, 25 

spissicornis, Lytta, 25, 27 

stigJTuita, Lytta, 150, 152 

stolida, Lytta, 3, 185, 186, 191 

stygica, Lytta: 2, 3, 4, 22, 46, 48, 
163, 185-192; larval hosts, 18; com- 
pared with chloris, 182, 183 

Stygica Group: 26, 161-163; food 
plants, 31; distribution, 32, 35; 
larvae, 158, phylogeny and syste- 
matics, 158-159; level of differen- 
tiation of species in, 182 

suavissima, Lytta. See Linsleya 

Subgenera: phylogeny of, 30-31 

sublaevis, Lytta: 22, 23, 35, 162, 163- 
165; compared with Meloe, 24; 
type of Poreospasta, 157 

Subspecies. See Races 

subviolacea, Lytta, 80, 81, 82 

sumptuosa, Lytta. See Prolytta 

Sunflower, 61, 90 

suturella, Lytta, 27 

Swarming behavior, 12, 238, 239 

Symphoricarpos, 13 

Synonymies, 5 

Syringa, 13 

szekessyi, Lytta, 25 

Tabasco, 72 

taliana, Lytta, 27 

Tamaulipas, 61 

tarsalis, Lytta, 229, 230, 234 

Tarweed, 127 



tenebrosa, Lytta: 2, 3, 4, 43, 44, 48, 
115, 116-119, 192; larval host, 18; 
ultimate larval instar, 20; typical 
race, 118; northern coastal race, 
118; Central Valley race, 118 

Tenebrosa Subgroup, 26, 51, 115-116 

Tennessee, 231, 239 

texana, Lytta, 229, 230, 234 

Texas, 59, 61, 88, 89, 94, 95, 110, 
145, 154, 231 

Thermopsis montanus, 209 

thibetana, Lytta, 26 

Thistle, 60 

Tlaxcala, 72, 96 

Toads: eating Meloidae, 11 

Transcaspian region, 31 

Tricrania, 8 

Tuberculosis: treated with canthari- 
din, 10 

ulkei, Lytta, 3, 120, 121 
Ultimate larval instar. See Larvae 
unguicularis, Lytta, 36, 225, 235, 

2.38-239 
Urinary disorders: treated with can- 

tharidin, 10 
usta, Lytta, 25, 26 
Utah, 59, 89, 103, 112, 130, 133, 168, 

203, 208, 211, 213, 214 

validicornis, Lytta, 25 

variabilis, Lytta: 2, 12, 40, 41, 42, 
63-67; courtship, 14-15; larval host, 
18 

Variabilis Group: 15, 26, 32, 52, 53, 
61-62, 102; distribution, 32; food 
plants, 50; phylogeny, 50-51 

Variabilis Subgroup, 26, 51, 62-63 

Variation: polymorphic, 65-67; indi- 
vidual, 84, 92, 96, 99-100, 106, 
147, 165, 171, 181, 182, 183, 189- 
191; 217; melanistic, 130. See also 
Geographic variation 

vellicata, Lytta, 25 

Veracruz, 72, 81, 84, 96, 146, 152, 
156 

Verbena, 169 

Verbenaceae, 169 

Verbesina encelioides, 90 

Vermont, 237 

Vertex: defined, 5 



INDEX 



295 



vesicatoria, Lytta: 23, 27; first instar 
larva, 2, 19; reared on Ceratina 
pollen, 2; feeding habits, 3; ver- 
nacular names, 9; as cantharides, 9; 
source of cantharidin, 9, 10; activ- 
ity period of adults, 11; longevity 
of adults, 12; food plants, 13; court- 
ship, 14; mating, 15; oviposition, 
16; larval instars, 16-17; larval host, 
18; grub stage, 19; coarctate larval 
instar, 20; ultimate larval instar, 20; 
pupal stage, 20; type of Lytta, 22; 
distribution, 25 

Vesicatoria Group, 27 

Viburnum lentago, 238 

Vicia: 171, 200, 212; americana, 209 

Viguiera dentata, 97 

Virginia, 231 

viridana, Lytta: 3, 32, 44, 48, 205, 



213-215; seasonal distribution, 11; 
geographic distribution, 214 
vulnerata, Lytta: 3, 36, 39, 114, 128; 
130-136; activity period of adults, 
11; distribution, 35; California race, 
135; interior race, 135 

Washington, 134, 189, 211 

Washington Territory, 121 

West Virginia, 231 

Wheat, 238 

Willow, 212, 234 

Wisconsin, 237 

Wyoming, 88, 203, 208, 214, 237 

Yucca, 130 

Zacatecas, 65, 96 
Zonitis, 11 




ss, george neville (1946) : American 
Species of Amelanchier. 23 pis. 14 maps. 
Vol. 20, No. 2. Cloth only, $2.00. 
kendeigh, s. Charles (1941): Territorial 
and Mating Behavior of the House Wren. 
32 figs. Vol. 18, No. 3. $1.50. 
( 1952) : Parental Care and Its Evo- 
lution in Birds. 35 figs. Vol. 22, Nos. 1-3. 
$4.00. 

kindred, james e. (1919): The Skull of 
Amiurus. 8 pis. Vol. 5, No. 1. $1.25. 

kramer, sol (1950): The Morphology 
and Phylogeny of Auchenorhynchous 
Homoptera (Insecta). 6 charts. 16 pis. Vol. 
20, No. 4. $2.00. 

kudo, richard roksabro (1934): Studies 
on Some Protozoan Parasites of Fishes of 
Illinois. 8 pis. Vol. 13, No. 1. $.75. 

(1944) : Morphology and Develop- 



ment of Nosema notabilis Kudo; Parasitic 
in Sphaerospora polymorpha Davis, a Para- 
site of Opsanus tau and O. beta. 12 pis. 7 
figs. Vol. 20, No. 1. $1.25. 

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Studies on Gregarines I— Including De 
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